Bull. nat. Hist. Mus. Lond. (Zool.) 63(1): 1-12

Issued 26 June 1997

A new species of Microgale (Insectivora,

Tenrecidae), with comments on the status of

four other taxa of shrew tenrecs or

PAULINA D. JENKINS __ !

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 SBD.

CHRISTOPHER J. RAXWORTHY

SE NE r mY,

Division of Herpetology, University of Michigan, Museum of Zoology, Ann Arbor, Michigan 48109, USA. Bball

RONALD A. NUSSBAUM

Jt JUL ivd/

PRESENTEC

Division of Herpetology, University of Michigan, Museum of Zoology, Ann Arbor, Michigan 48109, USA.

Synopsis. A new species of Microgale is described from rainforest localities in Madagascar. Evidence is presented that M.

drouhardi is a distinct species, with M. melanorrhachis as a synonym, and similarly that M. pulla is a synonym of M. parvula.

INTRODUCTION

The Family Tenrecidae is currently divided into four subfamilies,

three of which, the Geogalinae, Oryzorictinae and Tenrecinae are

endemic to Madagascar, while the fourth, the Potamogalinae, occurs

only inAfrica (see Hutterer, 1993). Within this highly diverse family,

the most diverse of all are the shrew tenrecs belonging to the

Oryzorictine genus Microgale Thomas, 1882, which at various

times has been subdivided into four genera or sub-genera, compris-

ing as many as 22 species. Despite the revision by MacPhee (1987)

based on specimens available at that time in museum collections, in

which only ten species were considered valid, the species composi-

tion of Microgale remains unclear. Subsequent to this revision, the

number of specimens of Microgale available has probably doubled

as a result of several recent expeditions to different localities in

Madagascar. These expeditions have attempted to provide an inven-

tory of the small mammal fauna, for taxonomic and biogeographic

purposes and for the development of conservation strategies. These

intensive surveys suggested that some of the species synonymised in

MacPhee’s revision, are in fact distinct (Nicoll & Rathbun, 1990;

Raxworthy & Nussbaum, 1994; Stephenson, 1995; Jenkins ef al,

1996; Goodman et al, 1996) and revealed the presence of several

undescribed species of Microgale (Jenkins, 1988, 1992, 1993; Jenkins

et al, 1996). Specimens from five widely separated localities, dis-

tinctive in external appearance, are believed to be conspecific and to

represent an additional undescribed species, the description of which

is given below.

During the course of these surveys, good samples of adult and

juvenile specimens of Microgale were collected from a wide range

of localities, allowing re-evaluation of the specific status of four

taxa, M. drouhardi Grandidier, 1934, M. parvula Grandidier, 1934,

M. melanorrhachis Morrison-Scott, 1948 and M. pulla Jenkins,

1988. The first three species were originally described from juvenile

specimens, a problematic situation in this genus, where the decidu-

ous and permanent dentitions may differ considerably, so causing

confusion over the correct specific attribution of juveniles and adults

(see MacPhee, 1987). Microgale drouhardi, M. parvula and M.

pulla were also known only from their respective type localities, and

M. parvula andM. pulla only from their holotypes. BothM. drouhardi

‘

a |

and M. melanorrhachis are of uncertain status and were considered

to be synonyms of M. cowani Thomas, 1882 by MacPhee (1987) but

evidence is presented below to show that M. drouhardi is a distinct

species with which M. melanorrhachis is synonymous. Likewise, it

is demonstrated that M. pulla is a synonym of M. parvula.

MATERIALS AND METHODS

In each of the surveys listed above, small mammals were collected

in pitfall traps, usually operating for five to ten days at each site. For

detailed information on collection methods see Raxworthy &

Nussbaum (1994; 1996) and Goodman et al (1996).

Measurements, in millimetres, were recorded using dial calipers

and a microscope measuring stage. External measurements include

head and body length (HB) from the tip of the nose to the

distalmost point of the body (at base of tail); tail length (TL)

measured from the base of the tail to the end of the distalmost

vertebra, excluding terminal hairs; hind foot length (HF) from the

heel to the distal part of the longest toe, excluding the claw; ear

length (EL) measured from the notch at the base of the ear to the

distalmost edge of the pinna; weight (WT) measured with Pesola

spring scales, animals weighing less than 10 gm within 0.2 gm and

10-100 gm within 0.5 gm. Cranial measurements were taken as

follows: condyloincisive length (CIL) cranial length from first

upper incisor to occipital condyle; upper toothrow length (UTL)

from anterior of first upper incisor to posterior of third upper

molar, parallel to the long axis of the skull; breadth of braincase

(BB) the greatest distance measured across the squamosals; height

of braincase (BH) greatest height in the midline from basioccipital

to parietal.

The dental nomenclature follows that of Mills (1966), Swindler

(1976), Butler & Greenwood (1979) and MacPhee (1987). Dental

notations are given in parentheses in the text; premaxillary and

maxillary teeth are denoted by upper case, mandibular teeth by

lower case, as follows: incisor (I/i), canine (C/c), premolar (P/p),

molar (M/m); a prefix “d’ indicates deciduous teeth, thus (dI) refers

to a deciduous upper incisor.

Age classes are defined as follows:

Infant: individuals in which the deciduous antemolar dentition

and the molars are not fully erupted; premaxillary, parietal and

basioccipital sutures unfused.

Juvenile: individuals in which the molars are fully erupted and the

deciduous antemolar dentition is erupted and in the process of

replacement by the permanent teeth; cranial sutures are in the

process of fusing. The eruption sequence of the permanent teeth

has been subdivided into four stages by MacPhee (1987).

Adult: individuals with a fully erupted permanent dentition; cra-

nial sutures generally fused although their position is usually

clearly marked.

Abbreviations used for institutions include, BMNH-— The Natural

History Museum, London [formerly British Museum (Natural His-

tory)]; FMNH — Field Museum of Natural History, Chicago; MCZ—

Museum of Comparative Zoology, Harvard; UMMZ-— University of

Michigan, Museum of Zoology, Michigan; USNM — National Mu-

seum of Natural History, Washington [formerly United States

National Museum].

Abbreviations used for protected sites in Madagascar: PN — Parc

National; RNI — Réserve Naturelle Intégral; RS — Réserve Spécial.

Other abbreviations used are: RAN — UMM field tag series; gm —

grams; km — kilometres; m — metres; mm — millimetres.

RESULTS

Microgale fotsifotsy sp. nov.

Figs 1-5

HOLOTYPE. UMMZ 168468 (RAN 38784) adult male, fixed in

formalin, preserved in alcohol, skull extracted. Collected by

Christopher Raxworthy [CR] 13 January 1992.

TYPE LOCALITY. Camp 2, Antomboka River Fitsahana, Parc Na-

tional de la Montagne d’Ambre, Antsiranana Fivondronana,

Antsiranana Province 12°29'S 49° 10'E, altitude 650m, rain forest.

P.D. JENKINS, C.J. RAXWORTHY AND R.A. NUSSBAUM

PARATYPES. Camp 2, Antomboka River Fitsahana, Parc National

de la Montagne d’ Ambre, Antsiranana Fivondronana, Antsiranana

Province 12°29'S 49° 10'E, altitude 650-670m, rain forest: UMMZ

171056 (RAN 38596), juvenile; BMNH 1996.278 (RAN 38648),

adult female; UMMZ 171057 (RAN 38710), juvenile; UMMZ

168470 (RAN 38740), juvenile; UMMZ 168466 (RAN 38752),

juvenile; UMMZ 171058 (RAN 38753), adult male; UMMZ 168467

(RAN 38754), adult male; UMMZ 168469 (RAN 38791), adult

male; UMMZ 171059 (RAN 38821), adult male. Collected by CR

2-17 January 1992. All specimens fixed in formalin, preserved in

alcohol, skulls extracted.

Camp 1, Antomboka River, Parc National de la Montagne d’ Ambre

12°32'S 49°10'E, altitude 1150m, rain forest: UMMZ 171055 (RAN

38192), adult female, collected by CR 23 November 1991. Fixed in

formalin, preserved in alcohol, skull extracted.

Parc National de la Montagne d’ Ambre, 5.5 km SW of Joffreville

[Ambohitra], 12°31'S 49°10'E, altitude 1000m, disturbed rain for-

est! FMNH 154590 adult male; FMNH 154591 juvenile male;

FMNH 154592 juvenile male; FMNH 154593 juvenile male; FMNH

154594 juvenile; FMNH 154595 juvenile male; FMNH 154596

juvenile male. Collected by Steven Goodman 28 March-1 April

1994. All skins and skulls.

REFERRED MATERIAL. RS d’Ambatovaky, Soanierana-Ivongo

Fivondronana, Toamasina Province 16°51'S 49°08'E, 600m, in rain

forest: BMNH 91.247.

RNI de Zahamena, Ambatondrazaka Fivondronana 17°40'S—

17°42'S 48°46'E, 850-1180m: UMMZ 171060-171065; BMNH

96.279.

Maitso, RNI d’Andringitra, 22°10'S 46°50'E, 1400 m, in dis-

turbed forest, 20 yards [18.3 m] from stream; found drowned in

puddle on top of shelter 5 feet [1.5 m] above the ground: BMNH

95.257. 40 km S of Ambalavao, along Volotsangana River, RNI

d’ Andringitra, 22°13'S 46°58'E, altitude 1210 m, in montane forest:

FMNH 151646-151647.

Marosohy Forest, 16 kmWNW of Ranomafana-Sud, Fivondronana

Tolagnaro [T6lanaro Fivondronana], Toliara Province, 24°34'S

46°48'E, 650m, in rain forest: USNM 578787; USNM 578887.

Fig. 1 Dorso-lateral view of Microgale fotsifotsy to show external features.

A NEW SPECIES OF MICROGALE

Table 1 Dimensions of adult Microgale fotsifotsy presented as range, mean + standard deviation and number of specimens in parentheses.

Combined PN Montagne d’ Ambre

650—1150m

Head and body length 63.9-81.0 63.9-70.0

69.0+5.11 66.2+2.17

(17) (8)

Tail length 71.4~-94.0 71.4-82.7

79.8+6.16 77.3+4.31

(16) (7)

Hind foot length 14-18 14-16

15.71.03 15+0.71

(17) (8)

Ear length 11-16 11-15

14.1+2.03 12.641.22

(16) (8)

Ratio of tail length 1.0-1.3 1.1-1.2

to head and body length 1.2+0.08 1.2+0.05

(16) (7)

Condyloincisive length 19.8-21.6 19.8-20.9

20.6+0.55 20.3+0.32

(16) (8)

Upper toothrow length 9.5-10.6 9.5—10.0

10.0+40.37 9.7+0.16

(16) (8)

Maxillary breadth from 5.8-6.6 5.8-6.3

M3-M3 6.1+0.26 6.0+0.17

(16) (8)

Braincase breadth 8.6-9.7 8.6-9.0

9.1+0.38 8.840.15

(16) (8)

Ratio of tail length 3.64.4 3.64.1

to condyloincisive 3.9+0.25 3.8+0.20

length (15) (7)

DIAGNOSIS. Digits of fore and hind feet and tail tip light coloured,

contrasting with darker coloration of head, body and tail. Ears pale

and conspicuous. Third upper and lower incisors (13 and i3) small, [3

slightly greater in crown height than distostyle of second upper

incisor (12), i3 subequal in height to posterior accessory cusp of

second lower incisor (i2); i2 greater in breadth than first lower

incisor (il); upper and lower canines greater in crown height than

second upper and lower premolars (P3 and p3)

DESCRIPTION. Small to medium in size, tail longer than head and

body (see Figs 1-2 and Table 1). Pinnae prominent and conspicu-

ous, pale in colour, reaching beyond eye if pressed forward along

head. Dorsal pelage soft in texture and grizzled yellowish brown

and grey; hairs with silvery grey bases, bright buff distally, usually

with dark brown tips; guard hairs slender, dark brown to black,

some with pale tips. Ventral pelage light grey with buff or reddish

wash; individual hairs with light silvery grey bases and light cream

tips. Tail more or less bicolored, grey brown above, light grey buff

below; with contrastingly light coloured tip, usually with thin pen-

cil of white hairs. Tail scale hairs moderately dense, approximately

three scales in length. Fore and hind feet brown with contrasting

light coloured digits, often with light lateral line along outer side

of foot. Fifth digit of hind foot elongated and scarcely shorter than

second digit; cheiridia on the hind feet elongated. Skull small to

medium in size (see Table 1). Rostrum moderately broad; interor-

bital region moderately short; maxillary process of zygoma at right

angles to long axis of cranium; braincase broad and short (see Fig.

3). Dentition illustrated in Figs 3-4. First upper incisor (I1) robust,

slightly proodont, distostyle prominent, lingual cingulum present,

buccal cingulum with accessory cusp; second upper incisor (12)

with small anterior accessory cusp, distostyle and prominent

RNI Zahamena RNI d’ Andringitra Marosohy forest

850-1180m 1200—1400m 650m

66.7-69.2 70.1-81.0 64, 77

68.2+1.02 76.7+5.8

(4) (3) (2)

75.0-84.0 85-94 71, 79

79.5+3.21 89.3+4.51

(4) (3) (2)

16-17 15-18 15, 16

16.5+0.50 16.341.25

(4) (3) (2)

15-16 16-17 12, 16

15.7+0.47 16.7+0.47

(3) (3) (2)

1.1-1.3 1.1-1.3 1.0, 1.1

1.2+0.07 1.2+0.09

(4) (3) (2)

20.7-21.6 20.7-21.2 19.8

21.1+0.38 21.0+0.29

(4) (3) (1)

10.2-10.5 10.0-10.6 O)S)

10.4+0.13 10.3+0.3

(4) (3) (1)

6.0-6.5 6.1-6.6 5.8

6.3+0.21 6.3+0.21

(4) (3) (1)

9.2-9.6 9.5-9.7 9.1

9.5+0.15 9.6+0.12

(4) (3) (1)

3.6-3.9 4.04.4 4.0

3.8+0.13 4.2+0.21

(4) (3) (1)

anterolingual cusp; [3 small, slightly greater than the distostyle of

12 in crown height, trace of distostyle evident in unworn teeth.

Upper canine (C) much greater than second upper premolar (P3) in

crown height, with indistinct anterior accessory cusp, distostyle

and lingual cingulum present. First upper premolar (P2) with small

anterior accessory cusp and distostyle; mesostyle of P3 small but

distinct, anterior ectostyle distinct and distostyle present;

mesostyle of third upper premolar (P4) large and distinct, anterior

ectostyle broad and distinct but posterior ectostyle and distostyle

distinct or indistinct; talon large and bicuspid in some specimens.

Talon of first and second upper molars (M1 and M2) large and

bicuspid; third upper molar (M3) subequal to or broader than M2.

Diastemata present between I1 and [2 and between I3 and C, C and

P3. Lower first incisor (i1) procumbent, with distinct hypoconulid;

lower second incisor (i2) spatulate, subequal in crown height but

broader than il and much larger than very reduced third lower

incisor (i3); 13 subequal in height to posterior accessory cusp of i2.

Lower canine (c) procumbent, anterolingual cingulum present but

anterior accessory cusp lacking. First lower premolar (p2) smaller

than c and anteroflexed, anterior cusp very small or absent in some

individuals. Protoconid of second lower premolar (p3) lower in

crown height than that of third lower premolar (p4) and paraconid

present; p4 and first and second lower molars (m1 and m2) lack

distinctive features, except for anterior and posterior buccal cin-

gula in most specimens. Talonid of third lower molar (m3)

incomplete and reduced to a hypoconulid, hypoconid indistinct or

absent, entoconid and entoconid ridge absent and entoconid basin

indistinct or absent. Diastemata generally absent, although present

in some individuals between c and p2.

Deciduous Dentition Deciduous anterior teeth smaller than per-

manent anterior teeth, in particular first upper and lower deciduous

4 P.D. JENKINS, C.J. RAXWORTHY AND R.A. NUSSBAUM

incisors (dI1 and dil) respectively considerably smaller than I1 and

il (see Figs 4 and 5). Principal cusp of dIl slender, distostyle

present, no anterior accessory cusp; morphology of deciduous sec-

ond upper incisor (dI2) similar to I2 but smaller; deciduous third

upper incisor (dI3) very small, less than height of distostyle of dI2.

Deciduous upper canine (dC) subequal in height to deciduous

second upper premolar (dP3), distostyle distinct, anterior accessory

cusp present. Trace of anterior cuspid on deciduous first upper

premolar (dP2); deciduous second upper premolar (dP3) with dis-

tinct mesostyle and distostyle present. Deciduous first lower incisor

(dil) subequal in height to second deciduous lower incisor (di2) but

considerably smaller in occlusal area; di2 similar in morphology to

i2, with or without a small lingual cusp. Deciduous third incisor

(di3) shed in all specimens examined. Deciduous lower canine (dc)

and first lower premolar (dp2) similar in morphology to respective

permanent counterparts but smaller, with trace of anterolingual

cuspid; deciduous second lower premolar (dp3) taller than de (un-

like permanent dentition), metaconid present or indicated.

Eruption sequence partially determined: 13 to [3, I1 to il, P2/p2,

12/i2, P4/p4; P3/p3 and C/c last teeth to erupt but sequence remains

to be determined.

INTRASPECIFIC VARIATION. The known populations of this

undescribed species are geographically widely separated and this is

reflected in the moderately high degree of intraspecific variation.

There is slight variation in pelage coloration: specimens from PN de

la Montagne d’Ambre are generally paler than those from RS

d’Ambatovaky, RNI de Zahamena and RNI d’Andringitra which

also often have a reddish buff wash; specimens from RNI de

Zahamena often show a reddish buff wash ventrally, while those

from RNI d’ Andringitra have a buff wash. Specimens from the RNI

d’Andringitra and RNI de Zahamena populations are larger on

average than either of the other populations (see Table 1), particu-

larly in cranial dimensions. The two specimens from Marosohy,

although geographically closer to the RNI d’ Andringitra specimens

in southeastern Madagascar are, however more similar to the PN de

Fig. 2 Dorsal view of skins from left to right of Microgale drouhardi la Montagne d’ Ambre specimens in size, as is the single juvenile

(FMNH 154565), M. fotsifotsy (FMNH 154590) and M. parvula specimen from RS d’ Ambatovaky. There is some evidence of corre-

(FMNH 151620). lation between size and altitude; specimens from localities at lower

Fig. 3. Cranium of Microgale fotsifotsy (FMNH 154590) from left to right, dorsal, ventral, and lateral view of cranium and mandible.

A NEW SPECIES OF MICROGALE

-———_—_—____4

Fig. 4 Buccal view of left permanent anterior dentition of Microgale

fotsifotsy (UMMZ 168468) I1—P2 above, 11—p2 below. Scale = | mm.

Fig.5 Deciduous right anterior dentition of Microgale fotsifotsy (FMNH

154529) buccal view of dI1—dP2 above; buccal view middle, lingual

view below of dil, di2, i3, dp2. Below right lingual view of right m3.

Scale = 1 mm.

Table 2 Altitudinal size variation in adult Microgale fotsifotsy.

Altitude Altitude

650m-670m 850m—1400m

Condyloincisive length 19.8—20.4 20.0-21.6

20.1+0.25 20.8+0.49

(7) (9)

Upper toothrow length 9.5-9.9 9.7-10.6

9.7+0.15 10.2+0.28

(7) (9)

Braincase breadth 8.5-9.1 9.0-9.7

8.9+0.16 9.5+0.23

(7) (8)

Ratio of tail length to 3.54.0 3.64.4

condyloincisive length 3.8+0.17 4.0+0.24

(6) (9)

altitudes averaging smaller than their counterparts from higher

altitudes (see Table 2).

ETYMOLOGY. Fotsifotsy is Malagasy for pale or whitish. The name

is used here as ‘the pale one’ in reference to the light coloured fore

and hind feet and tail tip, as well as the dorsal pelage which is paler

than in most other species of Microgale.

COMPARISONS WITH OTHER SPECIES. Microgale fotsifotsy differs

from other species of Microgale in a number of external and

craniodental features. The combination of light body coloration and

conspicuously pale coloured feet and tail tip, and the large, pale ears

are unique to this species (see Figs 1 and 2). The early stages of the

eruption sequence apparently differs from that ascribed toM. cowani

and M. pusilla Major, 1896b by MacPhee (1987) and toM. soricoides

Jenkins, 1993. In the sequence common to the three latter species, 13

and i3 erupt in the first stage, followed in sequence by P2, p2, p4, and

I] and il at the end of the second stage. In the specimens available

of M. fotsifotsy, the sequence progresses from i3 to I3, to I1 toil, to

p2 then P2, so that, unusually the first upper and lower permanent

incisors replace the deciduous teeth before any of the permanent

premolars have erupted. Microgale fotsifotsy resembles M.

longicaudata Thomas, 1882 in having elongated cheiridia and fifth

digits on the hindfeet and with a tail longer than head and body

length; the tail is, however, considerably longer in M. longicaudata

than in M. fotsifotsy (ratio of TL to HB 1.03—1.3, mean 1.2 + 0.08, n

= 16 in M. fotsifotsy; 1.7-2.4, mean 2.1 + 0.22, n = 11 in M.

longicaudata). Despite some similarities to M. longicaudata in

external proportions, the crania of the two species differ markedly.

That of M. fotsifotsy is larger and the braincase is relatively broader,

shorter and shallower than that of M. longicaudata in which the skull

is slightly concave in profile.

Despite the differences in size and external appearance, the skull

of M. fotsifotsy is more similar in overall proportions toM. soricoides

than to any other species. The skull of M. fotsifotsy is much smaller

and more delicate than that of M. soricoides. The rostrum is much

more slender and elongated but the maxillary processes of the

zygoma are flared at right angles to the long axis of the cranium in

both species, and the braincase is similarly short and broad, although

the occipital is proportionately much smaller inM. soricoides. There

are also more features common to the dentitions of these two species

than to others. In both M. fotsifotsy and M. soricoides, 11 is robust

and somewhat similar in its proportions, but that of M. fotsifotsy is

less pro-odont, and a larger cuspid is present on the buccal cingulum.

In both species, [2 is very similar in shape and proportions but [3 is

less reduced than in M. soricoides, in which P2 is also very reduced,

unlike M. fotsifotsy. These two species also differ considerably in the

anterior ectostyle of P3 which is distinct in M. fotsifotsy but very

reduced inM. soricoides. Inthe mandibular dentition of M. soricoides,

both i3 and p2 are very reduced and the latter is unusual in having a

single root; both of these teeth in M. fotsifotsy are reduced but less

markedly so and p2 has two roots. The lower canine is similar in both

species but p3 is more caniniform in M. soricoides. The third lower

molar shows a similar degree of reduction of the talonid in both

species.

The dentition of the new species is distinctive and does not readily

group with any of the species clusters described by MacPhee (1987)

but, as outlined above, does show some similarity to M. soricoides,

which may represent a separate cluster. The degree of development

of accessory cusps on I1-C is similar to that of members of the

cowani cluster but less marked than in the longicaudata cluster, also

the presence of interproximal gaps between the upper incisors

occurs in M. fotsifotsy and the cowani cluster. In contrast, M.

fotsifotsy is more similar to the /ongicaudata cluster in that the

buccal aspects of P3 and P4 are similar in shape, the lower canine

lacks a paraconid and the crown of p2 appears anteroflexed as inc,

due to the shorter convex anterior slope and longer concave posterior

slope. The relative proportions of the anterior teeth of M. fotsifotsy

differ from either the cowani or the longicaudata cluster, they are

similar to, but less extreme than those in M. soricoides, as shown in

Table 3.

Table 3 Relative proportions of the anterior teeth of Microgale fotsifotsy

in comparison with M. cowani cluster, M. longicaudata and M.

soricoides.

cowani cluster I1 >12>I3<C>P2< P3 C=P3

longicaudata Il>I2>I13<C>P2> P3 CSP3

fotsifotsy Il>I2»I13<C»P2< or= P3 (C5123)

soricoides Il » 12 » 13 « C »P2« P3 CS P3

cowani cluster il =i2>i3<c= or <p2<p3 c<p3

longicaudata il <12>i3<c> p2 < p3 c=p3

fotsifotsy il >i2»i3<c» p2 < p3 c2p3

soricoides il »12»i3«c» p2« p3 c=p3

Microgale drouhardi G. Grandidier, 1934

Microgale melanorrhachis Morrison-Scott, 1948

Microgale cowani Thomas: MacPhee, 1987, in part.

HOLOTYPE. MCZ 45034 (specimen A in original description)

juvenile female, body in alcohol, skull extracted, collected by

Monsieur Drouhard.

TYPELOCALITY. environs of Diego-Suarez [Antsiranana, c. 12°16'S

49°18'E-see MacPhee, 1987].

PARATYPES. MCZ 46007-46011 (specimens B-F in original de-

scription), MCZ 46012 (juvenile mentioned in original description),

all in alcohol. All from the same locality as the holotype.

REFERRED MATERIAL. environs of Diego-Suarez [Antsiranana, c.

12°16'S 49°18'E]: MCZ 46013-46019.

PN de la Montagne d’ Ambre, 5.5 km SW of Joffreville [Ambohitra],

12°31'S 49°10'E, 1000m, in relatively undisturbed rain forest: FMNH

154489; FMNH 154491; FMNH 154493-154494; FMNH 154499;

FMNH 154514—-154515; FMNH 154517; FMNH 154561-154564;

FMNH 154567-154571.

Camp 1, Antomboka River, PN de la Montagne d’ Ambre,

Antsiranana Fivondronana, Antsiranana Province, 12°32'S 49°10'E,

1150-1250m, rain forest: UMMZ 171000-171015; BMNH

1996.280; BMNH 1996.281.

5km S of Joffreville [Ambohitra], Mont[agne] d’Ambre, Diego

P.D. JENKINS, C.J. RAXWORTHY AND R.A. NUSSBAUM

Suarez [Antsiranana] Province, [c 12°32'S 49°10'E], rain forest:

USNM 341692; 6km S of Joffreville [Ambohitra], Mont[agne]

d’ Ambre, Diego Suarez [Antsiranana] Province, [c 12°32'S 49°10'E],

rain forest: USNM 341693.

Bekolosy, RS de Manongarivo, Ambanja Fivondronana, 14°03'S

48°18'E, 1150m, rain forest: UMMZ 171016—-171019.

Matsabory, RNI deTsaratanana, Ambanja Fivondronana, 14°09'S

48°58’'E, 2350m, rain forest: UMMZ 171020.

RS d’ Ambatovaky, Soanierana-Ivongo Fivondronana, Toamasina

Province 16°51'S 49°08'E-49°16'E, 360-600m: BMNH 91.220;

BMNH 91.221—91.226.

RNI de Zahamena, Ambatondrazaka Fivondronana 17°41'S—

17°42'S 48°46'E, 850-920m: UMMZ 171021—171027.

RNI de Zahamena, Vavatenina Fivondronana 17°44'S 48°59'E,

420-560m: UMMZ 171028-171039.

PN de Mantady, Moramanga Fivondronana, Toamasina Province,

18°51'S 48°27'E, 1100m, rain forest: UMMZ 168471-168474.

Didy, E of Lake Alaotra [Toamasina Province, c 18°02'S 48°32'E

— see Carleton & Schmidt, 1990]: USNM 328686.

Périnet [= Andasibe], near Moramanga, eastern Madagascar,

19°00'S 48°30'E, 3000 feet [915m]: BMNH 48.88 (holotype of

Microgale melanorrhachis).

43 km S of Ambalavao, junction of Sahanivoraky and Sahavatoy

Rivers, RNI d’ Andringitra, 22°13'S 47°00'E, 810m: FMNH 151627;

FMNH 151756. 45 km S of Ambalavao, E bank of Iantara River,

along Ambalamanenjana—Ambatamboay Trail, edge of RNI

d’ Andringitra, 22°13'S 47°01'E, 720m: FMNH 151626.

6 miles E of Ivohibe, 22°30' 47°00'E, 5000 feet [1525m], [high

rain forest]: BMNH 48.87.

TAXONOMIC NOTES. The original description of M. drouhardi was

based on several specimens, consisting of whole bodies preserved in

alcohol, with the exception of the holotype, which is a body with the

skull extracted. Grandidier (1934) believed that all but one of these

specimens were adult but MacPhee (1987) demonstrated that the

holotype is immature with the dentition partially deciduous (Stage 2,

see MacPhee, 1987: 13). As MacPhee emphasised, repeated

misidentifications and faulty taxonomic judgements have occurred

in this genus because of the problems of distinguishing between

deciduous and permanent dentitions. However, two of the speci-

mens mentioned in the type description are adult and, more critically,

another specimen (MCZ 46017) prepared as a skull and skeleton, is

adult and, although not mentioned in the original description, was

evidently available to Grandidier. Grandidier deposited his private

collection, including the original series of M. drouhardi, in the

Museum of Comparative Zoology, Harvard. In effect, this material

became unavailable to subsequent authors, such as Morrison-Scott

(1948) and Heim de Balsac (1972), who were unaware at which

institution in the the United States these specimens were held.

MacPhee was apparently the first author to re-examine this material,

he concluded that, although at the extreme of the size range, the

morphology of the teeth was such that the specimens were insepara-

ble from M. cowani and that M. drouhardi should be treated as a

synonym.

The type locality of ‘the environs of Diego-Suarez [Antsiranana]’

is unfortunately vague and, as pointed out by MacPhee (1987) and

Nicoll & Rathbun (1990), includes ecologically diverse habitats of

dry forest on limestone at Tendrombohitr’ Antsingy, Ankarana,

Analamera and Cap d’ Ambre, grassland and dry forest on the lower

slopes of the basaltic Montagne d’ Ambre, with rain forest at higher

altitudes of Montagne d’Ambre. Since most species of Microgale

are recorded from rain forest (M. pusilla Major, 1896b and M.

principula Thomas, 1918 contained in owl pellets of indeterminate

A NEW SPECIES OF MICROGALE

Table 4 Variation between populations of adult Microgale drouhardi. Dimensions given as range, mean+standard deviation and number of specimens in

parentheses.

PN Montagne d’ Ambre Bekolosy

1000—1250m 1150m

Head and body length 64-83 73.0, 73.3

75.8+5.39

(16) (2)

Tail length 67-83 70.5, 72.4

74.7+4.85

(16) (2)

Hind foot 16-19 17

length 17.6+0.77

(17) (2)

Ratio of tail length to 0.8-1.2 1.0

head and body length 1.0+40.10

(16) (2)

Condyloincisive length 22.5—23.9 22.0, 22.5

23.3+0.45

(18) (2)

Upper toothrow length 10.6—11.5 10.3, 10.7

11,140.21

(19) (2)

Maxillary breadth from —_6.0-6.7 6.2

M3-M3 6.4+0.14

(19) (2)

Braincase breadth 9.0-9.7 95.9.8

9.4+0.18

(19) (2)

Ratio of tail length tp 3.0-3.6 3.1, 3.3

condyloincisive length 3.2+0.21

(15) (2)

age, and M. brevicaudata Grandidier, 1899 are exceptions [for

discussion of these apparently anomalous drier habitats see MacPhee,

1987]) it is conjectured here that rain forest is also the most likely

source habitat for the type series of M. drouhardi. Additionally the

large size of these specimens suggests that they may have originated

from a high altitude locality (see section on variation), which is most

compatible with Montagne d’Ambre rather than any of the other

localities listed above.

In the original description, Grandidier described the dorsal pelage

as uniformly dark with the venter barely lighter in colour. The young

specimen mentioned in the type description (evidently an infant) is

described as having a black line along the mid-dorsum, which

according to Grandidier, disappears in adults. Re-examination of the

type series (by CR) shows that due to the discoloring and deleterious

effect of long-term storage in alcohol, these specimens are no longer

dark in colour but are now a dull reddish brown. Furthermore, a dark

dorsal stripe is discernable, which is obvious in some specimens,

where it is most evident on the head and anterior part of the body, but

faint and difficult to see in others. Shared features such as the

presence of a dorsal stripe, size and relative proportions of the body

and cranium, and morphology of the deciduous and permanent

dentitions, between the type series of M. drouhardi and specimens

recently collected (particularly from Montagne d’Ambre), lead to

the conclusion that they are conspecific.

Microgale melanorrhachis Morrison-Scott, 1948 was originally

distinguished by the presence of its dark mid-dorsal stripe. The

description was based on the skin and skull of the juvenile holotype

(believed to be adult by the author) from Perinet [Andasibe], plus a

damaged skin from another locality (Ivohibe). Following the origi-

nal description, an additional specimen was recorded by Eisenberg

& Gould (1970) from Didy, east of Lake Alaotra and references to

RS d’ Ambatovaky RNI Zahamena Didy PN Mantady

360-600m 420-1180m 1100-1150m

62.5-74.4 63.0-76.8 80.1 69.8-75.4

68.8+4.00 69.7+3.78 72.5+1.98

(7) (17) (1) (4)

52.9-62.3 52.7-62.1 - 63.7-71.0

56.7+3.24 58.3+2.83 67.7+2.66

(7) (16) (4)

13-15 14-15 - 16-18

14.1+0.64 14.6+0.49 17.0+0.71

(7) (17) (4)

0.8-0.9 0.8-0.9 0.9-1.0

0.8+0.05 0.9+0.05 0.9+0.04

(7) (15) (4)

21.0-21.6 20.5-22.3 D2 22.4, 22.5

21.340.22 21.4+0.45

(7) (17) (1) (2)

9.8-10.3 9.7-10.4 10.2 9.9-10.5

10.1+0.16 10.040.23 10.2+0.26

(7) (17) (1) (4)

6.1-6.3 5.9-6.4 6.2 6.3-6.9

6.2+0.09 6.1+0.17 6.6+0.11

(7) (17) (1) (4)

8.5-9.1 8.5-9.2 9.2 8.9-9.5

8.8+0.20 8.9+0.20 9.3+0.26

(7) (17) (1) (3)

2.4-2.9 2.6-2.9 ~ 3.0

2.7+0.17 2.740.12

(7)

(15)

(2)

the species were included in the literature (Heim de Balsac, 1972;

Genest & Petter, 1975). MacPhee (1987), however, observed that a

proportion of M. cowani also exhibited a variable tendency to a mid-

dorsal stripe, and he synonymised M. melanorrhachis with M.

cowani on the basis of its dentition. Subsequent authors disregarded

MacPhee’s opinion and continued to treat M. melanorrhachis as a

distinct species (Nicoll & Langrand, 1989; Nicoll & Rathbun, 1990;

Raxworthy & Nussbaum, 1994; Stephenson, 1995). The conclusion

drawn from the current study comparing recently collected large

series of adult and juvenile specimens from the localities listed

above with the type series of M. drouhardi and M. melanorrhachis,

is that they are conspecific.

DESCRIPTION. Medium sized (see Table 4), tail slightly shorter or

subequal to head and body length. Dorsal pelage dark brown, brown

or rufous brown with well demarcated, distinctly darker brown mid-

dorsal stripe, extending from crown of head to base of tail (see Fig.

2); hairs of dorsal pelage have light grey bases, red distally with

brown tips; hairs of mid-dorsal stripe dark brown for most of their

length, with grey bases, lacking red band; guard hairs also dark

brown with grey bases. Ventral pelage silvery buff, buff or rufous

buff, individual hairs having grey bases and buff or reddish buff tips;

transition between dorsal and ventral coloration moderately distinct.

Tail bicolored, dark brown above, buff or reddish buff below; tail

scales readily visible, scale hairs short, each overlapping c 2 scales

on basal third of tail. Hind feet dark brown on outer dorso-lateral and

ventral surface, reddish buff or buff on inner dorso-lateral surface.

Skull medium in size (see Table 4 and Fig. 6), with moderately

elongated and shallow but broad rostrum, nasals extend beyond

zygomatic plate into interorbital region; frontals slightly dorso-

laterally inflated; braincase moderately broad, short and shallow;

P.D. JENKINS, C.J. RAXWORTHY AND R.A. NUSSBAUM

Fig.6 Cranium of Microgale drouhardi (FMNH 154562) from left to right, dorsal, ventral, and lateral view of cranium and mandible.

parietal—supraoccipital junction rounded or subangular. Short dia-

stemata present between anterior teeth from I1 to P3 and usually

present on either side of p2; [3 often distinctively caniniform,

usually anteroflexed, anterior accessory cusp usually absent or very

reduced and distostyle may be reduced (see Fig. 7). Upper canine

long and slender, notably greater in crown height than other teeth in

upper dentition (including P3), root usually evident externally a

swelling in rostrum. Talonid of m3 slightly reduced, talonid basin

narrow and entoconid lacking (see Fig. 8).

VARIATION. There is a moderately high degree of variation in

pelage coloration between populations. Specimens from PN de la

Montagne d’ Ambre are usually dark brown, with or without a rufous

Fig. 7 Buccal view of left permanent anterior dentition of Microgale

drouhardi (FMNH 154514) 11—P2 above, i1—p2 below. Scale = 1 mm.

wash; those from Bekolosy, PN de Mantady, Didy and Andasibe,

also RS d’ Analamazaotra (see Stephenson, 1995) are bright rufous

or rufous brown dorsally, with a buff or rufous wash ventrally;

specimens from RS d’Ambatovaky are dull rufous brown dorsally,

grey ventrally; those from RNI de Zahamena are light rufous brown

dorsally, with a rufous wash ventrally; while specimens from RNI

Fig. 8 Deciduous left anterior dentition of Microgale drouhardi (UMMZ

168480): buccal view of dI1—dP2 above, buccal view of dil—dp2 below.

Below right lingual view of right m3. Scale = 1 mm.

A NEW SPECIES OF MICROGALE

Table 5 Altitudinal size variation in adult Microgale drouhardi.

Altitude Altitude

360m-920m 1000m—1250m

Condyloincisive length 20.5—22.3 21.1-23.9

21.4+0.47 2 23.0+0.78

(15) (23)

Upper toothrow length 9.7-10.5 9.9-11.4

10.1+0.23 10.8+0.47

(22) (25)

Braincase breadth 8.5-9.3 8.6-9.8

8.9+0.20 9.4+0.31

(22) (23)

Ratio of tail length to 2.5-2.9 2.7-3.6

condyloincisive length 2.7+0.14 3.1+0.21

(20) (20)

d’Andringitra (all juveniles) have a grey brown dorsum, grizzled

with yellowish buff. Although always present and distinctly demar-

cated, there is some variation in the width and length of the dorsal

stripe. The intensity of the rufous buff coloration of the venter,

undersurface of the tail and lateral surfaces of the hind feet also

shows within-population variability. The pelage of juveniles from all

localities is indistinguishable in coloration and degree of striping

from that of adults, which refutes the view of Grandidier (1934) that

striping is a juvenile characteristic and confirms MacPhee’s obser-

vation that striping occurs also in dental adults.

This species exhibits an exceptional degree of inter-population

size variation (see Table 4). Specimens from RS d’ Ambatovaky and

RNI de Zahamena are notably smaller than those from PN de la

Montagne d’ Ambre and Bekolosy, although the extremes shown by

these populations are bridged by specimens from PN de Mantady,

(and RS d’Analamazaotra, see Stephenson, 1995) which are inter-

mediate in size. Such a high degree of size variation is much greater

than encountered in any other species of Microgale so far investi-

gated. There is no apparent correlation between size and geographical

location, although there is some evidence of a link with altitude.

Specimens from lower altitudes are smaller, those from higher

altitudes average larger (see Table 5).

COMPARISONS. Microgale drouhardi is most similar in size and

dentition to M. cowani and M. taiva Major, 1896b. Microgale taiva

is another taxon regarded as a synonym of M. cowani by MacPhee

(1987), however recently collected specimens suggest that the two

species are distinct and the descriptions of both have been amplified

(see Jenkins ef al., 1996). Microgale drouhardi is readily distin-

guished from both species by the presence of the sharply demarcated,

dark, mid-dorsal stripe; also in life, M. drouhardi has a much paler

Table 6 Comparison of adult Microgale drouhardi, M. cowani and M.

taiva. Dimensions presented as range, mean+standard deviation and

number of specimens in parentheses.

M. drouhardi M. cowani M. taiva

Ratio of tail length to 2.4-3.6 2.5-3.1 3.54.2

condyloincisive length 3.0+0.27 2.9+0.19 3.9+0.19

(33) (14) (14)

Condyloincisive length 20.5—23.9 21.4-23.7 22.1—23.5

22.4+1.02 22.5+0.52 22.8+0.44

(39) (16) (14)

Ratio of anterior 41.6-50.9 52.8-56.4 49.1-52.8

dentition(I—P3) to 48.8+1.87 54.0+1.00 50.7+0.88

upper toothrow length (41) (16) (14)

Braincase breadth 8.5-9.8 9.8-10.4 9.6-10.3

9.2+0.36 10.1+0.19 9.9+0.22

(40) (18) (14)

venter than M. cowani and M. taiva. Microgale cowani is distin-

guished from M. drouhardi by the more elongated rostrum with

longer diastemata between the teeth of the anterior dentition, by the

broader, deeper braincase and by the presence of all elements of the

talonid of m3 (see Table 6). Microgale taiva has a relatively broader,

deeper braincase than that of M. drouhardi, the tail is relatively

longer, and the ratio of the anterior teeth (from I to P3) to the upper

toothrow fs greater on average (see Table 6).

Microgale parvula Grandidier, 1934

Microgale pulla Jenkins, 1988

HOLOTYPE. MCZ 45465, juvenile male, body in alcohol, skull

extracted, collected by Monsieur Drouhard.

TYPELOCALITY. environs of Diego-Suarez [Antsiranana, c. 12°16'S

49°18'E — see MacPhee, 1987]

REFERRED MATERIAL. Camp 1, Antomboka River, PN de la

Montagne d’ Ambre, Antsiranana Fivondronana, Antsiranana Prov-

ince, 12°32'S 49°10'E, 1125-1225m, rain forest! UMMZ

171043-171044.

Bekolosy, RS de Manongarivo, Ambanja Fivondronana,

Antsiranana Province, 14°03'S 48°18'E: BMNH 96.282.

RNI de Marojejy, Andapa Fivondronana, 14°26'S 49°46'E, 650—

900m: UMMZ 171045-171051.

Forét d’ Anandrivola, c 10 km southwest of Maintimbato Village,

c 40 km southwest of Maroantsetra, northeast Madagascar, 15°46'S

49°35'E, 450-625 m: BMNH 87.132 (holotype of M. pulla).

RNI de Zahamena, Ambatondrazaka Fivondronana 17°40'S

48°46'E, 1180-1270m: UMMZ 171052-171053.

RS d’Ambohitantely, Ankazobe Fivondronana 18°11'S 47°17'E:

UMMZ 171054.

PN de Mantady, Moramanga Fivondronana, Toamasina Prov-

ince, 18°51'S 48°28'E, 1100m: UMMZ 171040-171042; BMNH

96.283.

38 km S of Ambalavao, along Volotsangana River, RNI

d’Andringitra, 22°11'S 46°58'E, 1625 m: FMNH 151623; FMNH

151723; FMNH 151793-151794; FMNH 151801; FMNH

151805-151806. 40 km S of Ambalavao, along Volotsangana

River, RNI d’Andringitra, 22°13'S 46°58'E, 1210 m: FMNH

151722; FMNH 151764; FMNH 151766. 43km S of Ambalavao,

junction of Sahanivoraky and Sahavatoy Rivers, RNI

d’Andringitra, 22°13'S 47°00'E, 810 m: FMNH 151622. 45 km S

of Ambalavao, east bank of Iantara River, along Ambalaman-

enjana~Ambatamboay Trail, edge of RNI d’Andringitra, 22°13'S

47°01'E 720 m: FMNH 151621.

Ampamakiesiny Pass, Tolanaro Fivondronana, Toliara Province,

24°32'S 46°S1'E, 750-850m. UMMZ 167258; UMMZ 167261-

167263.

Marosohy Forest, 16 km WNW of Ranomafana-Sud,

Fivondronana Tolagnaro [T6lanaro Fivondronana], Toliara Prov-

ince, 24°34'S 46°48'E, 700-800m: USNM 578784-578785;

Marosohy Forest, near Tsitongatona River, 15 km WNW of

Ranomafana-Sud, Fivondronana Tolagnaro [Tolanaro Fivon-

dronana], Toliara Province, 24°34'S 46°48'E, 580m: USNM 578786.

Nahampoana, Vohimena Mountains; Tolanaro Fivondronana,

24°58'S 46°58'E, 120m: UMMZ 167233.

Manantantely, Vohimena Mountains, To6lanaro Fivondronana,

24°59'S 46°S8'E, 100-300m: UMMZ 167222-167223; UMMZ

167228-167229; UMMZ 167231.

TAXONOMIC NOTES. Microgale parvula was described originally

from a single specimen, erroneously considered to be adult by the

Table 7 Comparison of adult Microgale parvula and M. pusilla.

Dimensions presented as range, mean+standard deviation and number of

specimens in parentheses.

M. parvula M. pusilla

Head and body 49.6-64 52

length 56.1+3.8

(19) (1)

Tail length 46.5-66 72

54.8+5.9

(19) (1)

Hind foot length 9-11 12

10.0+0.64

(19) (1)

Ratio of tail length to 0.8-1.1 1.4

head and body length 1.0+0.08

(18) (1)

Condyloincisive length 15.5-17.1 15.6-16.8

16.4+0.36 16.4+0.4

(17) (6)

Upper toothrow length 6.9-7.8 7.2-1.9

7.4+0.21 7.6+0.19

(18) (9)

Maxillary breadth 43-4.6 4.9-5.2

from M3—M3 4.5+0.08 5.0+0.11

(19) (9)

Braincase breadth 6.4-6.9 6.8-7.3

6.7+0.15 7.0+0.16

(18) (6)

Braincase height 3.64.3 4.7-5.4

4.0+0.17 5.0+0.24

(18) (6)

Ratio of tail length to 2.9-4.0 4.4

condyloincisive length 3.4+0.33

(17) (1)

author but later demonstrated to have a deciduous dentition by

MacPhee (1987). Perhaps because its very small size presents

collection difficulties, no further specimens were recorded in the

literature nor represented in the major museum collections accessed

by MacPhee (1987). The dentition of this specimen was illustrated

by MacPhee (1987), who also corrected the measurements given in

P.D. JENKINS, C.J. RAXWORTHY AND R.A. NUSSBAUM

the original description. He was in no doubt about the validity of this

distinctively small species, which he grouped in the cowani cluster

on the basis of phenetic characters. The origin of the holotype of M.

parvula is the same as for M. drouhardi and similar arguments may

be applied, suggesting that it was most probably collected from

Montagne d’Ambre and from rain forest. It was assumed that the

species might be confined to northern Madagascar. Microgale pulla

Jenkins, 1988 was described from another single, although adult

specimen, from further south. MacPhee (personal communication)

advised that this specimen might simply represent the adult of M.

parvula and this possibility was mentioned in the original descrip-

tion of M. pulla, although the data available at that time suggested

otherwise. Recent collections from different localities extending

from Montagne d’ Ambre in the extreme north to the eastern forest as

far south as Manantantely, included adult and juvenile specimens

positively ascribable to the same taxon, directly associating the adult

pulla with the juvenile parvula. Microgale pulla was synonymised

with M. parvula (see Jenkins et al., 1996).

DESCRIPTION. Very small (see Table 7 and Fig. 2), tail subequal in

length to that of head and body. Dorsal pelage dark brown, ventral

pelage dark grey brown, tail and feet uniform dark grey brown.

Individual hairs of dorsal pelage with grey bases, orange red distally

with brown tips, guard hairs flattened, with grey bases and brown

tips. Hairs of ventral pelage with grey bases and brown or reddish

buff tips. Tail scales visible beneath moderately dense scale hairs,

2.5—3 scales in length. Skull very small, delicate and elongated in

appearance (see Fig. 9); rostrum slender, moderately short; brain-

case shallow and long, frontals and occipital large relative to parietals,

occipital condyles postero-dorsally orientated. Diastemata present

between I1 and [2 and on either side of C and P2; anterior and

posterior accessory cusps present on [2, I3 and P2. Diastema be-

tween c and p2. Talonid of m3 with well developed hypoconulid but

reduced hypoconid, entoconid and entoconid ridge, and narrow,

shallow talonid basin. See Fig. 10 for illustrations of permanent and

deciduous dentitions.

VARIATION. There is no obvious intraspecific variation in pelage

coloration or size between the populations studied, nor any evidence

Fig.9 Cranium of Microgale parvula (FMNH 151621) from left to right, dorsal, ventral, and lateral view of cranium and mandible.

A NEW SPECIES OF MICROGALE

of altitudinal variation. Variation may, however, be present but

obscured by the small sample size from individual localities.

COMPARISONS. Microgale pusilla Major, 1896b is the only species

with which M. parvula is likely to be confused because of similari-

ties in size. The two species differ in coloration: M. parvula is dark

grey brown dorsally, grading into grey ventrally, with a dark grey tail

which is only slightly paler ventrally: in contrast, M. pusilla is

reddish buffy brown dorsally, with a moderately abrupt transition to

the buffy grey brown venter and with the tail darker grey brown

above than below. Although the skull of both species is of a similar

length (see Table 6), the skull proportions differ considerably and, in

particular the braincase of M. parvula is shallower than that of M.

pusilla and the occipital condyles are more postero-dorsally aligned

(see Fig. 9 and Jenkins, 1988: Fig. 1). The major dental difference

between the two species is that p2 has a single root in M. pusilla but

is double rooted in M. parvula.

Fig. 10 Top: left permanent anterior dentition of Microgale parvula

(FMNH 151623) buccal view of I1—P2 above, buccal view of il—p2

below. Below: deciduous left anterior dentition of Microgale parvula,

buccal view of dI1—dP2 (FMNH 151806) above, buccal view of dil—dp2

(FMNH 151622) below. Lower right: lingual view of right m3 (FMNH

151622). Scale = 1 mm.

DISCUSSION

Purely on a phenetic basis, MacPhee (1987) grouped the species of

Microgale into six species ‘clusters’, as follows:

cowani Cluster: M. cowani, M. thomasi, M. parvula. Species

considered by MacPhee to be synonyms of M. cowani, but subse-

quently removed from synonymy, namely M. taiva (see Jenkins et

al., 1996) and M. drouhardi (this paper) continue to be assigned to

this group.

gracilis cluster: M. gracilis

longicaudata cluster: M. longicaudata, M. principula

pusilla cluster: M. pusilla

brevicaudata cluster: M. brevicaudata

dobsoni cluster: M. dobsoni, M. talazaci

Of those species described after MacPhee’s revision, Microgale

dryas Jenkins (1992) was considered to group with members of both

the cowani and gracilis clusters, while the unusual dentition of M.

soricoides Jenkins, 1993, was believed to be sufficiently distinct to

merit a separate cluster. Because of its greater resemblance in

craniodental morphology and proportions to M. soricoides rather

than any other species, M. fotsifotsy is placed in the soricoides

cluster.

Surprisingly, although Microgale fotsifotsy represents a new spe-

cies, the distribution of this shrew tenrec extends through almost the

entire length of the rain forest belt (from latitude 12°-24°S) in

Madagascar. It has been collected from six different regions:

Montagne d’ Ambre, Marojejy, Ambatovaky, Zahamena, Andringitra

and Marosohy Forest, near RNI Andohahela (Anosy Mountains).

The elevational range for this species is 600—1400m, suggesting that

it is primarily restricted to mid-altitude rain forest. Because trapping

in lower elevations at Ambatovoky, Zahamena and theAnosy Moun-

tains (as well as other sites) did not yield further examples of M.

fotsifotsy, we conclude that this species is absent from forest below

600m altitude.

The habits of this species remain largely unknown; most captures

so far have been made with pitfall traps in primary rain forest,

demonstrating that it is active on the forest floor. One specimen,

however, was found on top of a shelter 1.5m above the ground,

sug gesting that it may also be partly arboreal. This view is supported

by morphological features such as the relatively long tail and

elongated hind foot with a long fifth digit, adaptations which are

associated with arboreality in Soricidae (Hutterer, 1985) and occur

in a more extreme form in another species of shrew tenrec, M.

longicaudata, (see Thomas, 1918) a species for which there is some

behavioural evidence of arboreal adaptation (CR personal observa-

tion; Goodman, personal communication).

Microgale drouhardi and M. parvula are also widely distributed

in the rain forest regions of Madagascar. Both occur as far north

as Montagne d’Ambre (latitude 12°S) and as far south as

Andringitra (22°S) for M. drouhardi, and Manantantely, Vohimena

Mountains (25°S) for M. parvula. Microgale drouhardi has one of

the largest elevational ranges known for the genus, from 360-

2350m. The only other species with a similar elevational range is

M. talazaci Major, 1896a which occurs from 100m (Raxworthy,

unpublished) to about 2300m (Albignac, 1970). Microgale

drouhardi, M. talazaci (and M. cowani, Goodman personal com-

munication) appear to be the most montane Microgale known.

The elevational range of M. parvula is also large, between 100—

1550m altitude. None of these species of Microgale are specialists

of a single elevational zone, having been collected from low, mid

and even montane evergreen forest. Microgale drouhardi occurs

sympatrically in parts of its range with several other species simi-

lar in size and external morphology, notably M. cowani and M.

taiva. Obvious ecological differences have yet to be determined

between these species, except that M. drouhardi is known from

higher elevations. The only Microgale species similar in size to

M. parvula is M. pusilla, and based on MacPhee’s (1987) distribu-

tion summary, they are broadly sympatric over much of the eastern

rain forest belt (latitude 18°—21°S), although there is as yet no

evidence of microsympatry.

The broad elevational and latitudinal distributions of these rain

forest species of Microgale are surprising in as much as M.

parvula was previously only known by the holotype and M.

fotsifotsy was unknown. Conventional small mammal traps are

inefficient at trapping the smaller shrew-sized mammals, which

fail to trigger the trap mechanism. This may explain why M.

fotsifotsy, and more particularly, the very small, light weight, M.

parvula were previously unrecorded or rarely recorded. Microgale

drouhardi, although more commonly collected, is only represented

by small series in museum collections and, following the revision

of MacPhee (1987) has been considered as a synonym of M.

cowani. One of the main reasons for this situation is that most

Microgale material was collected in the last century (see MacPhee,

1987), with modern specimens being rare in museums. The pitfall

trapping method that we used in this study has proved to be

extremely productive at sampling Microgale populations.

ACKINOWLEDGEMENTS. We are greatly indebted to Jean Baptiste

Ramanamanjato, Achille Raselimanana, Angelien and Angeluc

Razafimanantsoa, for their assistance with this study. This research was made

possible through the cooperation of the Malagasy Ministére de l'Enseignement

Supérieur, the Ministére de la Production Animale et des Eaux et Forét, and

the Ministére de la Recherche Scientifique et Technique pour le

Développement.

We particularly thank Steven Goodman, Field Museum of Natural History,

Chicago for allowing us to examine recently collected material from RNI

d’ Andringitra and PN de la Montagne d’ Ambre, which included specimens

of the undescribed species, and also for bringing additional specimens of this

undescribed species in the collection of the USNM to our attention. For their

kind assistance and for the loan of specimens in their care, we are grateful to

William Stanley and John Phelps, Division of Mammalogy, Field Museum of

Natural History; Phillip Myers and Jim Birch, Division of Mammalogy and

Gregory Schneider, Division of Herpetology, University of Michigan, Mu-

seum of Zoology; Michael Carleton, Marc Frank and Helen Kafka, Division

of Mammalogy, National Museum of Natural History, Washington, D.C. For

two specimens we thank respectively, Tanya Barden and members of the

1993 Cambridge University expedition, Louise Ashmore, Juliet O’ Keefe,

Matthew Thomas and Oliver Tunstall Pedoe. Maria Rutzmoser, Museum of

Comparative Zoology, Harvard, kindly provided information on the holotypes

of M. parvula and M. drouhardi.

For their highly valued constructive comments and criticism of the manu-

script, we are indebted to Steven Goodman, Ross MacPhee (Department of

Mammalogy, American Museum of Natural History) and Darrell Siebert

(Department of Zoology, The Natural History Museum).

Photographs were prepared by Phillip Hurst, Photographic Unit, The

Natural History Museum.

This research was supported in part by grants from the National Science

Foundation (DEB 90-24505 and DEB 93-22600) and the National Geo-

graphic Society (5396-94). Logistic support was provided by the World Wide

Fund for Nature and Conservation International.

P.D. JENKINS, C.J. RAXWORTHY AND R.A. NUSSBAUM

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Bull. nat. Hist. Mus. Lond. (Zool.) 63(1): 13-26

Notes on the anatomy and relationships of

Sundasalanx Roberts (Teleostei, Clupeidae),

with descriptions of four new species from

Borneo

DARRELL J. SIEBERT |

Department of Zoology, Natural History Museum, Cromwell Road, London, U.K. SW7 SBD

CONTENTS

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SYNOPSIS. Sundasalanx has a prootic bulla and a recessus lateralis. Thus it is a clupeiform. The degree of caudal skeleton

consolidation found in Sundasalanx suggests relationship to the clupeid genus Jenkinsia. Sundasalanx is unusual among known

miniature clupeiforms. Its somatic development is equivalent to that of late stage unmetamorphosed larvae of other clupeiforms;

Sundasalanx is highly progenetic. Among fishes perhaps only Schindleria attains a comparable degree of progenesis. Sundasalanx

is widely distributed over Southeast Asia, with considerable species diversity. Descriptions are provided for four new Bornean

species: S. malleti Siebert and Crimmen; S. mesops Siebert and Crimmen; S. megalops Siebert and Crimmen; and S.

platyrhynchus Siebert and Crimmen.

Issued 26 June 1997

INTRODUCTION

Roberts (1981) described two new species of minute transparent

fishes, one from Khlong Falamee, Isthmus of Kra, southern Thai-

land, and the other from the Kapuas River, Kalimantan Barat,

Indonesia, as Sundasalanx praecox and Sundasalanx microps, te-

spectively. Sundasalanx is now known also from peninsular Malaysia

(Roberts, 1984; Kreutzman, 1993), Thailand north of the peninsula

(Roberts, 1984), and eastern Borneo north of the Kapuas River basin

(L. Parenti, pers. comm.). Four new species of Sundasalanx are

described below, three from the Barito River, Central Borneo,

Indonesia and one from the Kapuas river, West Borneo, Indonesia.

The new species of Sundasalanx appear to be larger than S. praecox

and S. microps but are very small fishes nevertheless, with observed

standard lengths not exceeding 30 mm.

Roberts (1981) considered Sundasalanx to be so unlike other

known fishes he erected a new family for it. While recognising its

unusual nature Roberts nevertheless felt it is allied with Salangidae

(icefishes), hence the name Sundasalanx, and its familial derivative,

Sundasalangidae. The following list of features was presented as

evidence in support of this conclusion: 1) jaw suspensorium consist-

ing of a single cartilaginous element; 2) 4° hypobranchial element

present; 3) pedunculate pectoral fins; 4)scaleless body; 5) absence of

symplectic; 6) absence of circumorbital bones; 7) myotomal muscles

not meeting at the ventral midline of the body; and 8) distal two-thirds

of maxilla curved beneath the head so that its ventral edge is directed

medially. Others have regarded Sundasalanx simply as a salangid,

arguing that if its relationship is with salangids then recognition of a

Sundasalangidae would render Salangidae paraphyletic (Fink, 1984;

Begle, 1991), if co-ordinate ranking is maintained in classification.

Johnson & Patterson (1996) recently have proposed a classification of

the Salmoniformes in which salangids are nested within the Osmeridae.

In their classification salangids are given lower rank. They plus

Mallotus make up the tribe Salangini.

All of the features enumerated above as suggestive of a relation-

ship with salangids, except possibly the curve of the posterior

portion of the maxilla to beneath the head, are simply aspects of a

physiognomy that is paedomorphic to an extreme. They are in fact

features suggestive of a larval stage of development and are a

consequence of a truncated ontogeny. As such they might not each

constitute independent evidence of relationship as each might be the

result of the same process or event that altered the development of

Sundasalanx. Furthermore, they amount to statements of absence,

which renders them ineffectual as evidence of relationship in the

absence of corroboration from other characters. Different evidence,

and a new radical hypothesis of relationship for Sundasalanx are

presented below.

MATERIALS AND METHODS

Comparative morphometry is presented with reference either to

Standard length (SL) or Head length (HL), each measured as

recommended in Hubbs & Lagler (1947). Head width (HW) was

taken as the width of the widest part of the head. Eye diameter was

measured from camera lucida tracings of heads. Counterstained

(C&S) materials were prepared following Dingerkus & Uhler (1977).

Since ossification in Sundasalanx is slight some materials were

cleared and then stained with alizarin only. Some very lightly

ossified elements were only apparent in material prepared this way.

Whether very lightly ossified elements were obscured by the blue

counterstain or dissolved by the acidic alcian blue solution during

the counterstaining process was not determined. Institutional abbre-

viations follow Leviton et al. (1985).

Anatomical notes are based on observations of cleared and stained

specimens: 17 Sundasalanx malleti sp. nov., from the Barito River;

10Sundasalanx mesops sp. nov., from the Barito River, 3 Sundasalanx

platyrhynchus sp. nov., from the Kapuas River; 2 paratypes of

Sundasalanx microps Roberts, from the Kapuas River, | paratype of

S. praecox Roberts, from peninsular Thailand; and | paratype of S.

megalops sp. nov., from the Barito River.

D.J. SIEBERT

MATERIAL EXAMINED. Materials of the new species are listed

below in the descriptions of new species.

Sundasalanx microps, paratypes, CAS 44220, 5 of 9 alc., 2 of 7

C&S, Indonesia, Kalimantan Barat, Kapuas R. basin, Kapuas R.

mainstream at Kampong Nibung, ca 100 km northeast of Selimbau,

5-6 Jul 1976, T.Roberts.

Sundasalanx praecox, paratypes, BMNH 1981.5.19:80-84, 4 alc.

ex., 1 C&S, Thailand, Isthmus of Kra, Khlong Falamee, a tributary

of Tale Sap, ca. 2 km W of Pak Payoon, 20 Jun 1970, T.Roberts.

NOTES ON THE ANATOMY OF SUNDASALANX

ROBERTS, 1981

Small size and light ossification make Sundasalanx difficult subjects

to study. Observations and interpretations that augment, or differ

from, those of Roberts (1981, 1984) and Kottelat (1991) are pre-

sented below. Although the relationships of Sundasalanx are

discussed later, comparisons are made here to clupeomorph,

clupeiform, or clupeid anatomy as an aid in interpreting Sundasalanx.

This is based on the conclusion that Sundasalanx is a clupeid, not a

salangoid.

Sundasalanx does not look like other juvenile or adult clupeids.

Rather, their physiognomy is very much like that of a late stage pre-

metamorphosis larva. Although there is a large literature concerning

the identification of larval clupeomorphs (McGowan & Berry, 1984)

surprisingly little has been written about their internal anatomy

(O’Connell, 1981).An exception is the gas bladder—inner ear—lateral

line system, of which detailed anatomical descriptions are available

for a number of clupeiforms (Allen et al., 1976; Blaxter & Hunter,

1982; O’Connell, 1981; Shardo, 1996).

Sundasalanx is distinctive. The peculiar structures Roberts (1981)

termed parapelvic cartilages (or bones; Fig. 1) are unknown else-

where among teleosts. Sundasalanx also possesses a highly derived

caudal skeleton (described below; Fig. 7A). A number of other

features listed by Roberts, mostly paedomorphic and not unique to

Sundasalanx, contribute to the generic diagnosis (Roberts, 1981).

The absence of hypobranchials 1-3 but with presence of hypo-

branchial 4, a cartilage bar uniting the shoulder girdle across the

ventral midline and a rayless pectoral fin supported by a single

cartilaginous plate rather than by separate radials are particularly

striking among the list.

Colour Pattern

Previous descriptions of Sundasalanx have been based on single

Fig. 1 Pelvic girdle of Sundasalanx malleti (anterior to left); A =

parapelvic bones, B = basipterygium, C = 1" pelvic fin-ray, D = radial,

scale bar = 0.5 mm.

RELATIONSHIPS OF SUNDASALANX

samples, albeit relatively large ones. Roberts (1984) suggested a

more detailed investigation of the colour pattern of different

populations of Sundasalanx was needed since every population

seemed to exhibit its own distinctive pattern, and implied that the

observed variation might be intra- rather than interspecific. Pre-

served materials studied here indicate that Sundasalanx species

share a basic colour pattern, but some species differ in certain

aspects and it varies with size and between sexes. Sex and size are

the two factors that appear to influence colour pattern most. Larger

specimens have more strongly developed colour patterns and males

have more vivid patterns than females or juveniles.

In life Sundasalanx is transparent except for the silvery cover

around the eyeballs, which decomposes to reveal black pigment

around the eye if specimens are conserved in formalin. Close

examination, however, of preserved specimens reveals an extensive

colour pattern (Figs 2,11,13-15). A small black spot is present

behind the eye in the prootic region (best seen in Fig. 14), associated

with the posterodorsal surface of what is herein identified as a

prootic bulla. This spot was observed in all species examined except

S. praecox, studied materials of which were largely depigmented.

Pigment deposition on the dorsal surface of the prootic bulla during

development has been observed in other clupeoids (Hoss & Blaxter,

1982), and is possibly a general feature of the group. Absence of

prootic bulla pigmentation in S. praecox is likely an artefact of

preservation.

A dark prepectoral mark, consisting of 1-5 melanophores, is

present just anterior to the pectoral fin in most individuals examined

(present in all large individuals; seen best in Figs 11, 13) of all

species except S. praecox. The extent of the mark appears to vary

interspecifically with the mark in Barito River species consistently

larger than that of species examined from elsewhere. In species with

a smaller mark it consists of one or two melanophores on the

dorsolateral surface of the connective tissue sheath surrounding the

sternohyoideus just anterior to the origin of the muscle on the

cleithrum. Species with a larger mark possess up to three more

melanophores on the rear wall of the gill chamber, dorsal to those on

the sternohyoideus.

Elements of a mid-ventral line of melanophores are present in all

species (Fig. 2). A midventral gular streak, dividing the sterno-

hyoideus muscle mass into right and left sides, is present in all

species. A midventral line of melanophores from behind the com-

partment in which the heart lies to the caudal fin, as described for

southern peninsular Malaysia materials by Roberts (1984), is present

in all species except S. praecox. In the region of the body cavity the

melanophores are associated with the ventral finfold or its remnants.

In the region of the anal fin, ventral midline pigmentation consists of

melanophores located between each anal-fin pterygiophore, one per

interpterygiophore space (best seen in Fig. 15); behind the anal fin

the line is continued as a midventral series of melanophores situated

between the ventral ends of myotomes of the caudal peduncle

myotomes.

A line of melanophores is present on each side of the body at the

ventral edge of myotomes in the region of the body cavity in all

species examined (Figs 11, 13-15). The line begins immediately

behind the shoulder girdle and ends at the anus, with the frequency

of melanophores usually greater than one per myotome, but less than

two per myotome. The line is continuous except in the region

immediately dorsal to the pelvic fins, from which melanophores are

absent.

In all species examined, a dark spot consisting of a single melano-

phore is located just anterior to the insertion of the pelvic fins (Figs

2, 11, 13-15), below the course of the myotomal line. An additional

spot is located near the anterior end of the basipterygium of S.

Fig. 2 Schematic representation of the melanophore pattern of

Sundasalanx in ventral view (anterior to left); A = gular streak of

midventral line, B = midventral line of the ventral fin fold, C =

myotomal line, D = melanophores of cleithral mark associated with the

sternohyoideus, E = cardiac compartment melanophores (present only in

males of S. microps, S. mesops, S. megalops, and S. platyrhynchus), F =

basipterygium melanophore (present in all species examined).

praecox. Males of S. malleti with mature, or maturing, testes show

additional pigmentation of the pelvic-fin girdle, with a ring of

melanophores encircling the basipterygium.

The lower caudal-fin lobe is marked by a broad swath of

melanophores in all species except possibly S. praecox. This lower

caudal-fin lobe swath is especially intense in materials identifiable

as male (Fig. 14). A few scattered melanophores were observed on

the upper caudal-fin lobe of a few individuals.

The posterior half of the compartment in which the heart lies is

marked with melanophores (Fig. 2) in males of S. microps, S.

mesops, S. megalops, and S. platyrhynchus, even in quite small

individuals. Examination of larger specimens in which it is obvious

that maturation of the testes has begun shows the mark is present

only in males; specimens with maturing or mature ovaries lack the

mark, as do all of the smallest specimens. The juxtaposition of the

anterior end of the myotomal lines to the pigmentation of the cardiac

compartment creates a very distinctive M-shaped ventral mark just

posterior to the pectoral girdle in males of those species with cardiac

compartment pigmentation (Fig. 2).

Soft anatomy

Gut

The gut of Sundasalanx is straight, with three sections clearly

distinguishable under gross microscopical examination of cleared

and stained specimens (Fig. 3). Length of gut measures approxi-

mately 62% of SL in the largest cleared and stained specimen

examined (24 mm SL). The first section, approximately 20% of total

gut length, is characterised by longitudinal folds, continuous through-

out its length. No obvious distinction between oesophagus and

stomach is apparent, although the posterior part of this section is

somewhat greater in diameter than the anterior part. The pneumatic

duct exits from the dorsal side of the posterior half of the stomachal

Fig. 3 Gut and gas bladder of Sundasalanx malleti (anterior to left); A =

anterior section of gut (oesophagus/stomach), B = middle section of gut

(anterior part of intestine), C = parapelvic bones, D = middle section of

gut (posterior part of intestine), E = posterior section of gut, F = gas

bladder, G = pneumatic duct, scale bar = 3 mm.

region. The second section, consisting of the intestine, is separated

from the oesophagus—stomach by an obvious sphincter, and meas-

ures approximately 73% of total gut length. The anterior third of the

intestine is marked by irregular folds, which are more vertically

oriented and regular in the region of the air bladder and parapelvic

bones. Posterior to the air bladder the folds are regular, nearly

vertical, and wavy. They appear to encircle the gut so that the

posterior half of the intestine looks very much like the external

appearance of an earthworm. The end of the intestine is conical and

projects slightly into the third section. The third section of the gut is

short, less than 10% of total gut length and is slightly greater in

diameter than the intestine. It also is characterised by wavy, vertical

folds, but these are more numerous (distinctly closer together) and

more wavy than those of the intestine.

Gonads

The gonads lie dorsal to the gut and when mature occupy nearly the

entire length of the body cavity, as described for S. praecox (Roberts,

1981). Testes exhibit the vertical striations characteristic of at least

some other clupeomorphs (Roberts, 1981; Whitehead & Teugels,

1985). Judging from enlargement of posterior sections of testes in

some individuals, maturation of testes proceeds from posterior to

anterior. Ovaries contain eggs of distinct size classes, egg size

presumably being an indication of degree of egg maturation. One

female contained 18 very large eggs, nine in each ovary arranged in

a single file. These eggs were slightly wider than the width of two

myotomes (nearly 1 mm in diameter) and occupied nearly the entire

length of the ovaries. Smaller eggs of two distinct size classes were

interspersed among the larger eggs (Fig. 4). Among clupeomorphs

the presence of distinct size classes of eggs in ovaries is considered

indicative of repeat spawning within a single breeding season.

Sundasalanx may thus have considerable fecundity even though

relatively few mature ova are present (can fit) within an ovary.

Roberts (1981) reported egg size in S. praecox of only 0.20-0.25

mm in diameter. Sundasalanx praecox may have smaller ova than

other Sundasalanx species, but among clupeiforms that are repeat

spawners final enlargement of the egg does not occur until shortly

before spawning (Blaxter & Hunter, 1982). Thus the difference in

egg size between Barito River species of Sundasalanx observed here

and S. praecox may be due to the stage of egg development at which

materials were preserved.

BSTOEOO

eee wy CG

Fig. 4 Outline drawing of the left ovary of Sundasalanx mesops (anterior

to left); A = large ovum, B = smaller size classes of ova, C = duct to

genital opening, scale bar = 2 mm.

Gas Bladder and Associated Structures

The gas bladder is located above the gut in the region of the pelvic

fins, and is somewhat constricted by the parapelvic bones (Fig. 3).

The pneumatic duct appears to enter the anterior end of the gas

bladder, an entry position unusual among clupeomorphs (White-

head and Blaxter, 1989). The gas bladder was deflated in most

specimens but in a few the anterior portion was partially filled with

gas. Nocturnal, facultative filling of the gas bladder by swallowing

air is well known among larval clupeomorphs. Thus no special

significance is attached to the observation that in some Sundasalanx

specimens the gas bladder is partially filled with air whereas in most

D.J. SIEBERT

Fig. 5 Photograph of the head of a skeletal preparation of Sundasalanx

malleti (BMNH 1996.7.18.15) showing the relative position, size, and

shape of the prootic bulla (A).

itis deflated. A duct leading from the gas bladder to the region of the

anus, usually present in clupeomorphs, was not observed.

A very large, vertically oriented, spindle-shaped bulla is present

in the prootic (Fig. 5). It is approximately 40% of head depth in

height, protrudes ventrally into the branchial chamber, and extends

dorsally nearly to the dorsal surface of the head. It is capped with

melanophores and is among the few structures other than the eyes

that can be seen on living Sundasalanx. A canal for the precoelomic

gas duct is apparent but the duct itself was not detected. Ossification

of the prootic bulla is not apparent in counter stained preparation,

only in preparations stained solely with alizarin.

Lateral to the prootic bulla lies a cavity, identified as a recessus

lateralis, on which the supraorbital, temporal, and infraorbital

laterosensory canals converge (Fig. 6). A small, circular fenestra on

the dorsolateral side of the prootic bulla communicates with the

Fig. 6 Schematic representation of the recessus lateralis and cephalic

laterosensory canals of Sundasalanx malleti (anterior to left); A =

temporal canal, B = infraorbital canal, C = supraorbital canal, D =

infraorbital bone, E = recessus lateralis chamber, F = fenestra on the

medial wall of the recessus lateralis, G = prootic bulla.

RELATIONSHIPS OF SUNDASALANX

recessus lateralis. A large fenestra to the rear of the recessus

lateralis communicates with the perilymphatic system.

Laterosensory canals

No laterosensory canals were observed other than the infraorbital,

supraorbital, and temporal canals which converge on the recessus

lateralis (Fig. 6). All are short The supraorbital canal does not

extend to even over the eye. The temporal canal does not reach the

shoulder girdle or form a supratemporal commisure with its counter-

part from the opposite side. The infraorbital canal does not extend to

beneath the eye.

Osteology

No part of the skeleton of Sundasalanx is heavily ossified. Ossifica-

tion of dermal elements appears to be exceptionally slight and many,

usually present in other teleosts, may be absent. For example, no

parietals or dermal ethmoid element were detected.

Caudal skeleton

The caudal skeleton of Sundasalanx is highly consolidated, and

somewhat reduced (Fig. 7A). Preural centrum 1 and ural centra 1

and 2 are consolidated into a single compound element, to which

uroneural | is apparently fused. Uroneural 2 is free, and in Barito

River Sundasalanx greatly expanded. Only a single epural is present.

The parhypural and hypural | are fused (observed in an ontogenetic

series of cleared and stained materials); the compound parhypural-

hypural is autogenous; hypural 2 is fused to the compound centrum;

hypural 3 is autogenous; hypural 4 is expanded to about the size of

F+G

hypural 1; hypural 5 is present. A hypural 6 was not observed. Bases

of caudal-fin rays appear unmodified.

Pelvic girdle

The two vertically oriented bones in the pelvic girdle of Sundasalanx

Roberts (1981) termed parapelvic cartilages are unknown elsewhere

among fishes. The anterior bone is stouter, more vertically oriented,

with its ventral end closely associated with the basipterygium and

first pelvic ray (Fig. 1). The posterior bone is more slender, usually

longer than the anterior bone, and is usually inclined forward, often

as much as 30° from vertical. It is not as closely associated with the

basipterygium as the anterior bone; there is often a considerable gap

between the ventral end of the posterior bone and the basipterygium.

The greater length and position of the posterior bone relative to the

basipterygium results in it extending above the dorsal end of the

anterior bone by as much as half its length. The length of the

posterior parapelvic bone appears to vary among species. In some

specimens of S. praecox the posterior bone is nearly twice the length

of the anterior bone whereas in specimen of S. megalops examined

the bones are subequal.

The origin of parapelvic bones is unknown. In some fishes pleural

ribs are connected to the pelvic girdle via ligaments and it would not

be unreasonable to suggest parapelvic bones might be modified ribs,

or ossifications of ligaments associated with the pelvic girdle.

Parapelvic bones are similar to pleural ribs in two respects. They are

preformed in cartilage (preformation of ribs in cartilage is wide-

spread among clupeoids; Patterson & Johnson, 1995), and they lie

internal to body musculature, not in myosepta. However, peculiar

Fig. 7 A) Caudal skeleton of Sundasalanx malleti (BMNH 1996.7.18.15), scale = .3 mm; B) Caudal skeleton of Jenkinsia (BMNH 1962.7.21.48-—5S0),

scale = .45 mm; A = compound centrum (PU1+U1+U2), B = epural, C = uroneural 2, D = hypural 5, E = hypural 4, F = hypural 1, G = parhypural

cartilage chevrons, unique among lower teleosts, are associated with

the lateral tips of epicentral intermuscular bones in many clupeoids

and the derivation of parapelvic bones from these structures should

also be considered.

Vertebral column

Vertebrae of Sundasalanx are lightly ossified, with neural arches

present on all vertebrae preceding the compound caudal vertebra.

Caudal vertebrae appear to ossify first. All neural arches preform in

cartilage and the anterior few may not ossify. The first neural arch

has a broader base than the others and neural spines do not appear to

form on the first 4 or 5 arches. No supraneurals are present. Haemal

arches also preform in cartilage. Cartilaginous basiventrals are

present on all preural centra, including the first. No ribs are present.

Intermusculars

No intermuscular bones or ligaments (Patterson & Johnson, 1995)

are present.

Dorsal- and anal-fin supports.

Middle radials do not develop in dorsal- and anal-fin pterygiophores.

Distal radials are clearly present. Proximal elements (=proximal +

middle radials) of the last few dorsal pterygiophores appear to fuse.

Jaws

Toothed premaxillary and maxillary bones make up the upper jaw.

No supramaxillary bones were detected. Dentary, anguloarticular,

and retroarticular bones were identified in the lower jaw.

Gill arches

The primitive complement of structures that make up the median

series of hyoid- and gill arch elements of teleosts, not all of which

may be present in adult forms, develops from three copulae (copula

1-3; Nelson, 1969). Derivatives of all three copulae are present in

Sundasalanx. The series includes (Fig. 8): the basihyal (derived

from copula 1); basibranchial 1-2 and basibranchial 3 (derivatives of

copula 2); and basibranchial 4-5 (derived from copula 3). Gill arches

1 and 2 articulate with basibranchial 1-2, gill arch 3 articulates with

basibranchial 3, and gill arches 4 and 5 articulate with basibranchial

4-5. Hypobranchials are not evident in the first three arches. Roberts

(1981) suggested they were fused to the median elements but such

fusion of lateral endoskeletal elements to median endoskeletal ele-

ments is rare (Nelson, 1969). A more likely explanation is that

hypobranchials 1—3 never condense from the hypo/ceratobranchial

precursors and thus never develop at all. Hypobranchial 4 is present

in all species examined, and a curious cartilage nodule, located in the

ligament between ceratobranchial 4 and 5, was seen in some speci-

mens of S. praecox, S. malleti, S. mesops, and S. megalops, but not

in S. microps or S. platyrhynchus, both from the Kapuas River. Five

conventional ceratobranchials are present. Four epibranchials and

three pharyngobranchials are present (Fig. 9). Epibranchials 24 are

short. Epibranchial | is large, occupying the space of pharyngo-

branchial 1 in addition to its own. Epibranchial 4 curves

anteromesially to articulate with the lateral posterodorsal corner of

pharyngobranchial 4, dorsal to the tooth plate associated with the

posteroventral side of pharyngobranchial 4. The efferent arterial

canal of epibranchial 4 is completely closed, with no sign of any line

of fusion of interbranchial 4. A small levator process projects

dorsally from the top of the ring of cartilage that forms this canal.

The levator exturnus IV, which has its origin on the posterodorsal

corner of the skull, inserts on this process. Epibranchial 3 is short,

and articulates with the posterolateral end of a large pharyngo-

branchial 3. The mesioposterior end of pharyngobranchial

D.J. SIEBERT

G Cc

Fig. 8 Lower branchial arches of Sundasalanx malleti (anterior to left); A

= ceratobranchial 1-5, B = hypobranchial 4, C = basibranchial 1-2, D =

basibranchial 3, E = anterior and posterior ceratohyals, F = hypohyal, G

= basihyal, scale bar = 1 mm.

Fig. 9 Dorsal view of left side upper gill arch elements of Sundasalanx

malleti (anterior to left); A = epibranchial 1, B = epibranchial 4, C =

tooth plates (circles indicate tooth positions diagramatically), D =

pharyngobranchial 2-4, scale bar = 0.25 mm.

3 articulates broadly with the anterior end of pharyngobranchial 4.A

large tooth plate is associated with the ventral surface of the poste-

rior half of pharyngobranchial 3.The uncinate process of epibranchial

3 is modified to form a canal for efferent artery 3. The artery is

completely ringed inS. praecox, so that the lateral end of epibranchial

3 looks similar to the lateral end of epibranchial 4. It is almost

completely ringed inS. platyrhynchus, but only partially encircled in

S. malleti, S. mesops, and S. megalops. Epibranchial 2 is also short,

but with a long uncinate process that articulates with the lateral edge

of pharyngobranchial 3, the point of articulation on pharyngo-

branchial 3 drawn out to form an ‘uncinate’ process in S. malleti.

Pharyngobranchial 2 is much longer than wide, extending from the

medial edge of epibranchial 2 in an anteromedial direction to the tip

of pharyngobranchial 3. No tooth plate was found to be associated

with pharyngobranchial 2. Epibranchial | is narrow near the articu-

lation with ceratobranchial 1, broadly spatulate at its medial edge. It

possesses a long uncinate process that articulates with the uncinate

process of pharyngobranchial 2. A separate pharyngobranchial 1

was not observed. Enlargement of epibranchial 1 may be the result

of fusion between it and pharyngobranchial 1, or of failure of

condensation of pharyngobranchial | from the epi-pharyngobranchial

anlagen in a shortened developmental programme.

Very thin perichondral ossification of ceratobranchial 5 was the

only ossification detected for endoskeletal gill arch elements. If

other elements are ossified their ossification is beyond the limit of

RELATIONSHIPS OF SUNDASALANX

Fig. 10 Schematic illustration of the opercular and branchiostegal series

of Sundasalanx malleti (anterior to left); A = opercle, B = subopercle, C

= interopercle, D = branchiostegals, E = preopercle, F = infraorbital

bone.

resolution of the staining procedures used, or the materials prepared

as whole mount preparations.

Dentition

Roberts (1981, 1984) reported teeth on the premaxilla and maxilla,

and that teeth are embedded directly in cartilage on the lower jaw

(Meckel’s cartilage), ceratobranchial 5, and pharyngobranchial 4 of

S. praecox and S. microps. Kottelat (1991) reported a pharyngo-

branchial 4 tooth plate in the species here named S. platyrhynchus,

rather than teeth embedded directly in cartilage. Materials prepared

only as alizarin preparations in this study indicate the conclusion

that some teeth in Sundasalanx are directly embedded into cartilage

is erroneous, and most likely due to examination of specimens

prepared by counter-staining for cartilage and bone.

In addition to those structures listed above, all species of

Sundasalanx examined were found to also have a tooth plate and

teeth associated with the underside of the posteromedial part of

pharyngobranchial 3. The size of the tooth plate appears to vary

among species. Sundasalanx praecox was found also to possess

teeth on the posterior end of basibranchial 1-2) in the floor of the

mouth, and small ‘vomerine’ tooth patches were found also in the

roof of the mouth, one on either side of the anterior tip of the

parasphenoid. The largest cleared and stained specimen of S. malleti

appears to have a tooth plate (toothless) associated with the posterior

end of the basihyal.

Suspensorium

The suspensorium is weakly ossified and not well differentiated.

Independent quadrate, metapterygoid, symplectic, or hyomandibula

ossifications were not detected in any species of Sundasalanx. The

posterior articulation of the suspensorium with the skull is broad. An

independent cartilaginous palatine was observed in all examined

species except S. praecox, the smallest species, in which a thin strut

of cartilage communicates between the palatine and the hyomandi-

bular complex. An extremely thin, sheet-like ossification under the

eye between the palatine and hyomandibular is here identified as the

mesopteryogoid.

Lower hyoid arch

The basihyal is a large expansive structure that essentially fills the

floor of the mouth anterior to the hyoid arch A cartilaginous

hypohyal is present, but apparently without differentiation into

upper and lower hypohyoid elements. Ossified anterior and poste-

rior ceratohyals are present, as is an unossified interhyal. Two, three,

or four branchiostegals, subequal in size, were found to be associ-

ated with the ventrolateral side of the posterior ceratohyal. The

number of branchiostegals present is not constant in any species of

Sundasalanx examined. The anteriormost branchiostegal is posi-

tioned at the anterior edge of the posterior ceratohyal, partially

overlapping the gap between the posterior- and anterior ceratohyal.

Succeeding branchiostegals are positioned posteriorly along the

ceratohyal and there is a distinct gap between the branchiostegal

series and the interopercle, which is associated with the posterola-

teral side of the posterior ceratohyal near the articulation of the

interhyal. The interopercle is distinctly larger than any member of

the branchiostegal series (Fig. 10).

Opercular series

An opercle, subopercle, interopercule, and preopercle were found in

all species examined (Fig. 10). All elements are very lightly ossified

and are easily overlooked, especially the preopercle. The opercle,

subopercle and interopercle form an overlapping series, with the

ventroanterior corner of the opercle lying external to the posterior

edge of the subopercle and the anterior edge of the subopercle lying

external to the posterior edge of the interopercle. The interopercle is

associated with the posterolateral side of the posterior ceratohyal,

near the interhyal. The preopercle is an elongate element. Its anterior

end lies lateral to the undifferentiated quadrate. More posteriorly it

lies lateral to the ventral portion of the cartilaginous structure that is

the undifferinated hyomandibula. Its posterior portion occupies the

space between the infraorbital and the opercle. No lateral sensory

canal was observed to be associated with the preopercle.

Infraorbital series

A single infraorbital element was detected. It is an extremely thin

ossification located in the interior of the bend of where the infraorbital

laterosensory canal turns in an anterior direction. The infraorbital

canal emerges from the recessus lateralis well behind the eye.

Consequently the sole infraorbital is positioned well posterior to the

eye.

Skull Roofing Bones

The only roofing bone identified is an ossification associated with

the supraorbital laterosensory canal and recessus lateralis. The bone

is very weakly ossified and difficult to detect. The pair, one on each

side, are here identified as frontals, because of the association

between the supraorbital laterosensory canal, the recessus lateralis

and the frontal in clupeiforms. Parietals, and dermal ethmoid ele-

ments were not detected.

RELATIONSHIPS

In the discussion that follows, the informal term salangid refers to

icefishes (=Salangidae of Roberts, 1984). This does not imply

criticism of the rank ascribed to them by Johnson and Patterson

(1996). Features identified as suggesting a relationship between

Sundasalanx and salangids (Roberts 1981, 1984; listed previously

[p. 5]) are all paedomorphic, with the possible expection of the

orientation of the posterior part of the maxilla. If maxilla orientation

in other larval clupeoids is found to be like that Sundasalanx then

this feature also is paedomorphic. The features are plesiomorphic

too, being features found in larvae of lower teleosts, and in larvae of

some euteleosts. Such a list, no matter how long, is nothing more

than an appeal to symplesiomorphy as evidence for the hypothesis of

relationship. However, as Roberts (1984) indicated the question

remains as to whether the apomorphic condition of neoteny in

Sundasalanx and salangids is a synapomorphy or a homoplasy.

Derived, non-paedomorphic, features unique to both Sundasalanx

and salangids can corroborate a Sundasalanx-salangid relationship,

thereby establishing neoteny as synapomorphic for Sundasalanx

and salangids.

The recent re-examination of lower eutelostean relationships by

Johnson & Patterson (1996) provides a context for the search for

non-paedomorphic features that might confirm a Sundasalanx—

salangid relationship. The relevant groups and characters (numbered

sequentially) are: euteleosts — 1) supraneurals develop caudally, 2)

uroneural 1 with a membranous outgrowth (stegural), 3) caudal

median cartilages present; salmoniforms — 4) epicentrals with carti-

lage rods distally, 5) epicentral and epineural intramuscular bones

lacking proximal forks; salmonoids + osmeroids — 6) derm- and

supraethmoid separate, 7) postemporal fossa open, 8) a single

supramaxilla, 9) upper pharyngeal tooth plate 4 absent, 10) anterior

epineurals not fused to neural arches; 11) epipleurals absent, 12)

distal parts of posterior neural and haemal arches forming a keel, 13)

uroneural 2 anterodorsal to uroneural 1, 14) scales without radii, 15)

nuptial tubercles present, 16) diadromy; osmeroids — 17) orbit-

osphenoid absent, 18) basisphenoid absent, 19) articular reduced,

20) gill rakers toothless, 21) preural centrum 1 and ural centrum 1

fused, 22) cleithrum with a narrow columnar process toward cora-

coid, 23) no postcleithrum; osmerids — 24) short hyomandibular

crest, 25) opercular with anterodorsal notch, 26) levator process

present on epibranchial 4, 27) uroneural | fused to preural centrum

1, 28) caudal scutes absent, 29) extrascapular fragmented into

several ossicles, 30) posterior dorsal pterygiophores fused, 31)

adipose cartilage present, 32) egg with adhesive membrane;

osmerines — 33) otic bulla (=saccular recess) somewhat inflated, 34)

keel formed by posterior neural and haemal spines absent; salangins

— 35 ethmoid endoskeleton long and unossified, 36) 1st pectoral

radial unmodified, 37) 4th pectoral radial multifid distally, 38) males

with modified anal fin endoskeleton; and Icefishes (=salangids) —

39) anterior margin of metapterygoid above quadrate, 40) antorbital

bone absent, 41) 1 supraneural present, 42) 4th pectoral radial

articulates with glenoid, 43) dermal plate absent from basibranchials.

At least 19 of these characters are ‘absence characters’, or ‘revers-

als’ to a more primitive condition, leaving just 24 as ‘presence

characters’ .As with Roberts’ list, resemblance between Sundasalanx

and salangids due to the first class characters requires confirmation

from congruence with the second class of characters. Character 32,

egg with adhesive membrane, was not checked in this study. Of the

remaining 24 characters from the second class of characters

Sundasalanx can be shown to have only four, fusion of preural

centrum | with ural centrum | (21), levator process on epibranchial

4 present (26), uroneural 1 fused to preural centrum | (27), and

posterior dorsal pterygiophores fused (30). Sundasalanx simply

lacks the rest, either because they are primitively absent or because

Sundasalanx is so underdeveloped they never appear in its ontogeny.

All of the four that are present in Sundasalanx are also present

among clupeocephalan, or even elopocephalan, fishes. The evidence

for a close relationship among Sundasalanx and icefishes should be

regarded as non-existent.

Sundasalanx is highly paedomorphic and as obvious from com-

parison to the above lists establishing its relationship presents

certain difficulties. It is not, however, wholly paedomorphic and the

non-paedomorphic features of Sundasalanx suggest a relationship

not with salangids, nor any other euteleostean, but with clupeiforms,

and further to dussumieriine clupeids.

Prootic bullae are found only among clupeomorph fishes and the

recessus lateralis is found only among clupeiforms-(Greenwood et

D.J. SIEBERT

al, 1966; Grande, 1985). Sundasalanx is thus a clupeiform. Estab-

lishing the relationships of Sundasalanx within Clupeiformes is less

easy. Many external and internal features used to elucidate relation-

ships within the order (Grande, 1985) are absent; all scute and scale

characters widely used in the identification of clupeiforms are

absent from Sundasalanx, which is completely scaleless and lacks

ribs and supraneurals. However, Sundasalanx exhibits a derived,

highly consolidated, caudal skeleton. Derived features are: 1) fusion

of preural centrum 1 (PU1), ural centrum 1 (U1), ural centrum 2

(U2), and the first uroneural into a single element; 2) reduction of the

number of epurals to 1; 3) reduction of the number of hypurals to 5;

4) fusion of the parhypural and hypural 1; 5) an expanded hypural 4;

and 6) absence of the extensions of the middle caudal fin rays

characteristic of clupeiforms. Among clupeiforms only the

spratelloidin dussumieriine genus Jenkinsia, a marine, Caribbean

endemic, approaches the degree of caudal skeleton consolidation

found in Sundasalanx (Fig. 7B). The pertinent modifications are

(Grande, 1985): 1) fusion of PUI, U1, and U2 into a single element;

2) reduction of the number of epurals to one; and 3) expansion of

hypural 4 to a size equal to that of hypural 1. Fusion of PU] with U1

is also known in some pellonulines, some engraulids and Clupeonella,

but these lack the other derived caudal skeleton features of

spratelloidins (Grande, 1985). Sundasalanx is thus a spratelloidin,

and possibly the sister-group of Jenkinsia.

DISTINCTIVENESS OF SUNDASALANX

PRAECOX

The discovery of materials seemingly intermediate in eye size

between. praecox and S. microps led Roberts (1984) to suggest that

distinction between them at the species level needed further consid-

eration, with the implication that there might be only one widespread

species of Sundasalanx. Sundasalanx praecox and S. microps were

re-examined for this study. I conclude that Roberts’ (1981) original

assessment of the specific status of S. praecox is correct. It is

different from other species of Sundasalanx in so many details that

it stands out as the most distinctive of all the described species. No

other known species of Sundasalanx possesses any palatal or

basibranchial teeth. It has far fewer vertebrae, many more upper jaw

teeth, larger pharyngeal tooth plates with many more teeth, larger

and more numerous gill rakers, a relatively longer posterior parapelvic

bone than any species known from Kalimantan, and lacks a midventral

line of melanophores associated with the ventral fin-fold.

DESCRIPTIONS OF NEW SPECIES

Sundasalanx malleti Siebert and Crimmen, sp. nov.

(Fig. 11)

HOLOTYPE. MZB 6096, 26.4 mm SL, Indonesia, Kalimantan

Tengah, Barito River basin, Sungai Barito at Muara Laung, dip nets

and seines, 20-22 Feb 1991, D. Siebert, A. Tjakrawidjaja, O.

Crimmen, and A. Effendi.

PARATYPES. MZB 6097 (20), collection data as for holotype.

BMNH 1996.7.18.147—311 (164), collection data as for holotype.

USNM 320689 (5), collection data as for holotype.

REFERRED MATERIAL. BMNH 1996.7.18.315—324 (10), C&S, In-

donesia, Kalimantan Tengah, Barito River drainage, sand bars of

RELATIONSHIPS OF SUNDASALANX

Table 1 Selected mensural characters of Sundasalanx malleti, S. mesops, S. megalops, and S. platyrhynchus; the mean + standard deviation is followed by

the sample minimum — maximum; sample size is reported in parentheses if different from that reported under each species’ name.

S. malleti n = 10

Eye (% SL) 4.2+0.3 3.8-— 4.7 5.2+0.3

Eye (% HL) PIS UB 19.9-23.2 25553

Snt (% SL) 4.3+0.4 3.5- 4.8 (9) 3.8+0.4

Snt (% HL) 2U221.7 18.4-23.9 (9) 18.6+1.7

Snt:eye 1.0+0.1 0.8— 1.2 (9) 0.7+0.1

Head d. (% SL) 8.2+0.8 6.3— 9.0 7.9+0.5

Head |. (% SL) 20.0+0.8 18.8-21.1 20.7+0.7

Head w. (& SL) 9.7+0.7 8.6-10.6 (9) 10.5+0.4

Table 2 Number of vertebrae of species of Sundasalanx; counts for S.

praecox and S. microps are from Roberts (1981). Analysis of variance

with a post hoc test of location reveals S. praecox to be significantly

different from S. microps, S. platyrhynchus, S. malleti, and S. mesops at

p<0.001.

Vertebrae

37 38 39 40 4] 42 43 44

S. praecox 2 7

S. microps

S. platyrhynchus

S. malleti

S. mesops

S. megalops

— oon — Ye

Table 3 Dorsal- and anal-fin branched ray counts for species of

Sundasalanx. Sundasalanx praecox, §. microps, S. platyrhynchus, S.

malleti, and S. mesops were tested for difference in number of dorsal-

and anal-fin branched rays with one-way analysis of variance, with

Tukey’s post hoc HSD test to locate differences of means. Sundasalanx

platyrhynchus differs from S. mesops in number of branched dorsal-fin

rays at p < 0.01; for branched anal-fin rays S. microps and S.

platyrhynchus test as significantly different from S. praecox, S. malleti,

and S. mesops with p < 0.01.

D-fin rays A-fin rays

Se Oi. NO) petite ie sn f14 dain 16,417

S. praecox 4 | | a” ate.

S. microps FM me i 2 2 aay

S. platyrhynchus 3° 15) 14 | rh Sy Dea

S. malleti 4 16 23 3 OT 1629135 rou

S. mesops a 26° 8 Ss 23" 9 2

2 I I

S. megalops

Fig. 11 Holotype of Sundasalanx malleti, female. MZB 6096, 26.4 mm

SL.

S. mesops n = 30

S. megalops n = 2 S. platyrhynchus n = 20

4.5— 5.7 5.9— 6.0 5.140.4 4.2—5.7 (18)

21.4-27.5 27.5—29.0 23.8+1.2 21.7-25.9 (16)

3.1— 4.8 3.5—4.9 4.040.7 2.9- 5.2

15.6—22.9 17.3-22.6 18.141.9 15.3-21.4 (19)

0.6— 1.0 0.6— 0.8 0.8+0.1 0.6- 1.0 (18)

7.3- 9.1 9.4-9.6 8.340.7 7.2— 9.5

18.7-21.8 20.4—21.6 21.4+1.2 19.6-23.2 (18)

9.8-11.2 9.4 (1) 10.940.5 10.2-12.0 (21)

Sungai Murung around Project Barito Ulu basecamp on Sungai

Murung, seine, 12 Feb 1991, D. Siebert, O Crimmen, and A.

Tjakrawidjaja.

DIAGNOSIS. A photograph of S. malleti is presented as Fig. 11;

selected morphometrics are summarised inTable | and vertebral and

fin-ray counts are summarised in Table 2 and Table 3. It is a species

of Sundasalanx with a relatively small eye (3.8-4.7% SL, 19.7-

23.2% HL); snout long (18.4—23.9% HL); and with snout:eye ratio

usually greater than | (0.8—1.2). Prepectoral blotch extensive; ma-

turing or mature males (large individuals) with basipterygium ringed

by melanophores; no melanophores on posterior wall of cardiac

compartment; lower caudal-fin lobe with a broad swath of

melanophores. Vertebral number 40-44 (median=42); dorsal-fin

rays 8—11 (median= 10); anal-fin rays 13-17 (median=14). Premax-

illa with 4—5 teeth, maxilla with 15-17 teeth; dentary with 10-12

teeth; Sth ceratobranchial tooth plate large, with 5—6 teeth in princi-

pal posterior row, 2 or 3 rows of smaller teeth anterior to principal

row; 3rd pharyngobranchial tooth plate large, longer than wide,

subequal to or larger than 4th pharyngobranchial tooth plate; 4th

pharyngobranchial tooth plate large, nearly 1/2 size of 4th pharyngo-

branchial, with principal row of larger teeth along posterior edge.

Sundasalanx malleti is easily distinguished from its Barito River

eye

1.0

10 15 20 25 30

Fig. 12 Scatter plot of Eye versus Standard length for Barito River

species of Sundasalanx, with linear smoothing function superimposed

on points for S. malleti and S. mesops; A= S. megalops, O = S. mesops,

x = S. malleti.

Fig. 13. Holotype of Sundasalanx mesops, female, MZB 6098, 20.7 mm

SE:

Fig. 14 Holotype of Sundasalanx megalops, MZB 6100, male, 25.5 mm

SL. The melanophore cap of the prootic bulla is clearly visible, the

myotomal pigment line is clearly interrupted above the pelvic fins,

melanophores associated with anal fin pterygiophores are clearly

visible, and the vivid swatch of melanophores on the lower caudal fin

lobe characteristic of males is clearly evident. The wavy striations of the

posterior half of the intestine are discernible posterior to the pelvic fins.

Fig. 15 Holotype of Sundasalanx platyrhynchus, MZB 5944, 19.2 mm

SL.

congeners and S. platyrhynchus by size of eye, snout length, and

colour pattern. Other Barito River Sundasalanx have discernibly

larger eyes (Figs 11-15; Table 1), have a snout:eye ratio < 1, lack a

ring of melanophores around the basipterygium in-maturing males,

D.J. SIEBERT

Fig. 16 Dorsal view of the head of Sundasalanx platyrhynchus.

and possess melanophores on the posterior wall of the cardiac

compartment in males. Table 4 summarises diagnostic features of all

described species of Sundasalanx.

SEXUAL DIMORPHISM. Sundasalanx malleti is sexually dimorphic

for colour pattern. Unbleached cleared-and-stained materials that

are identifiable as male or as female show a difference between sexes

in pigmentation of the basipterygium and lower caudal-fin lobe.

Males have a more intense mark on the lower caudal-fin lobe, the

result of a greater density of melanophores that make up the lower

caudal lobe swath. Males with maturing or mature testes also

possess additional pigmentation around the basipterygium that was

not observed in any female. Minimally, females, males, and juve-

niles possess a single melanophore just anterior to the pelvic-fin

origin. The basipterygium of males is marked by additional

melanophores along its side and across its posterior edge between

the pelvic-fin bases so that it is completely encircled with

melanophores.

ECOLOGY. Sundasalanx malleti was captured in turbid flowing

waters over silty, clayey substrates along banks adjacent to main-

stream currents and at depths up to 1 m. Children with fly-screen

scoop nets, working bankside shallows before dusk for small fishes

for the evening meal, captured them in great abundance. The same

shallows worked during daylight usually yielded no or few

Sundasalanx. Thus S. malleti probably inhabits deeper waters dur-

ing daylight hours, moving to shallower areas at dusk, possibly to

escape predation. Fishes captured in the same habitat with S. malleti

were many species of small and juvenile cyprinids and catfishes,

Nemacheilus sp., Homaloptera sp., and chandids.

Remains of insects and crustaceans are observable in guts of

cleared and stained specimens.

ETYMOLOGY. This species is named after John Valentine Granville

Mallet, former Prime Warden of the Worshipful Company of Fish-

mongers, whose enthusiasm, encouragement, and support have

made possible the continuation of a research programme on fresh-

water fishes of Southeast Asia.

RELATIONSHIPS OF SUNDASALANX

Sundasalanx mesops Siebert and Crimmen, sp.nov.

(Fig. 13)

HOLOTYPE. MZB 6098, 20.7 mm SL, Indonesia, Kalimantan

Tengah, Barito River drainage, Sungai Laung at desa Maruwei, O°

21.986'S, 114° 44.103'E, miscellaneous catches, 15—18 Jul 1992, D.

Siebert, A. Tjakrawidjaja and O. Crimmen.

PARATYPES. MZB 6099 (10), collection data as for holotype.

BMNH 1996.7.18.2—51 (50), collection data as for holotype.

REFERRED MATERIAL. BMNH 1996.7.18.52—56 (5), Indonesia,

Kalimantan Tengah, Barito River drainage, small tributary of Sungai

Maruwei approx. | km upstream from desa Maruwei, O° 21.986'S,

114° 44.103'E, rotenone, 15 Jul 1992, D. Siebert, A. Tjakrawidjaja

and O. Crimmen. BMNH 1996.7.18.57—66 (10), C&S, collection

data as for BMNH 1996.7.18.52-56. BMNH 1996.7.18.67—109

(43), collection data as for BMNH 1996.7.18.52-56. BMNH

1996.7.18.110-146 (37), Indonesia, Kalimantan Tengah, Barito

River drainage, Sungai Barito approx. 2 km below Muara Laung,

beach seine at dusk, 8 Jul 1992, D. Siebert, A. Tjakrawidjaja and O.

Crimmen. BMNH 1996.7.18.312 (1), 26.5 mm, Indonesia,

Kalimantan Tengah, Sungai Barito drainage, sand bank of Sungai

Joloi upstream from Sungai Busang, seine, 7 Feb 1991, D. Siebert,

A. Tjakrawidjaja and O. Crimmen. BMNH 1996.7.18.313-314 (2),

26.0—29.5 mm, Indonesia, Kalimantan Tengah, Barito River drain-

age, sand bars of Sungai Joloi upstream from Sungai Busang, seine,

8 Feb 1991, D. Siebert, O. Crimmen, A. Tjakrawidjaja.

DIAGNOsIS. A photograph of S. mesops is presented as Fig. 13;

selected morphometrics are summarised inTable | and vertebral and

fin-ray counts are summarised in Table 2 and Table 3. It is a species

of Sundasalanx with a relatively large eye (4.5-5.7% SL, 21.4—

27.5% HL); and relatively short snout (15.6—22.9% HL). Vertebral

number 40-41 (median=41); dorsal-fin rays 8—10 (median=9); and

anal-fin rays 13—16 (median=14). Prepectoral mark well developed;

posterior wall of cardiac compartment with melanophores;

basipterygium without ring of melanophores. Premaxilla with 3—5

teeth; maxilla with 16—18 teeth; dentary with 12-13 teeth in single

row; 5th ceratobranchial tooth plate with 3-5 teeth in principal

posterior row, teeth anterior to principal row small and few; 3rd

pharyngobranchial tooth plate small, with 3 or 4 teeth; 4th pharyngo-

branchial tooth plate relatively small, about 1/3 size of 4th pharyngo-

branchial, with principal row of larger teeth along posterior edge.

Sundasalanx mesops is easily distinguished from its Barito River

congeners by size of eye, colour pattern, and upper pharyngeal

dentition. The eye of S. mailleti is smaller; the eye of S. megalops is

larger (Figs 11—14). Colour pattern features that distinguish it from

S. malleti are listed above in the diagnosis for S. malleti. Sundasalanx

malleti has larger upper pharyngeal tooth plates, with more teeth; S.

megalops possesses fewer teeth in jaws and smaller upper pharyn-

geal tooth plates with fewer teeth (see below). Sundasalanx mesops

can usually be distinguished from S. platyrhynchus of the Kapuas

River, West Kalimantan by dorsal- and anal-fin ray counts (Table 3).

SEXUAL DIMORPHISM. Cleared and stained materials in which

male and female specimens are clearly identifiable reveal Sunda-

salanx mesops is sexually dimorphic for colour pattern. Females

differ from males in pigmentation of the cardiac compartment and

lower caudal-fin lobe. Males have a more intense mark on the lower

caudal-fin lobe; the lower caudal-fin lobe of the female specimen

with the largest eggs is not as dark as that of males with only partially

mature testes. Males with maturing or mature testes also possess

pigmentation of the posterior wall of the cardiac compartment which

was not observed in any female.

ECOLOGY. Sundasalanx mesops was captured downstream from

creek mouths in shallows over sandy or silty substrates during

daylight hours. During a spate specimens were also taken from

inundated grassy stream banks.

DISTRIBUTION. Sundasalanx mesops was taken from several lo-

calities throughout the upper Barito River basin, Central Kalimantan,

Indonesia, from the widest variety of habitat sizes. On present

evidence it has the widest distribution of Barito River Sundasalanx.

ETYMOLOGY. The name mesops, is a combination of meso, Greek

for middle, and ops, Greek for eye. It is in reference to the size of its

eye relative to other Barito River Sundasalanx species.

Sundasalanx megalops Siebert and Crimmen, sp. nov.

(Fig. 14)

HOLOTYPE. MZB 6100, 25.5 mm SL, Indonesia, Kalimantan

Tengah, Sungai Barito drainage, sand and silt bank at the mouth of

Sungai Sapen, a small left hand tributary of Sungai Joloi above

Sungai Busang, seine, 6 Feb 1991, D. Siebert and O. Crimmen.

PARATYPE. BMNH 1996.7.18.1; (1), 25.0 mm, C&S, collection

data as for holotype.

DIAGNOSIS. A photograph of S. megalops is presented as Fig. 14;

selected morphometrics are summarised in Table | and vertebral and

fin-ray counts are summarised in Table 2 and Table 3. It is a species

of Sundasalanx with a deep head (approximately 9.5% SL); large

eye (approximately 6.0 per cent SL, 28% HL); and short snout

(snout:eye ratio < 1). Prepectoral mark well developed; posterior

wall of cardiac compartment with melanophores; basipterygium

without ring of melanophores. Meristics of paratype as follows:

vertebral number 41; dorsal-fin rays 10, anal-fin rays 14-15. Pre-

maxilla with 2—3 teeth; maxilla with 12—14 teeth; dentary with 8-10

teeth, arranged in 2 rows in rear portion of dental arcade;

ceratobranchial 5 tooth plate with 5—6 teeth in principal posterior

row, teeth anterior to principal row small and few; pharyngo-

branchial 3 tooth plate small, with only 2 or 3 teeth; pharyngo-

branchial 4 tooth plate small, about 1/3 size of pharyngobranchial 4,

with principal row of larger teeth along posterior edge.

Sundasalanx megalops is easily distinguished from its Barito

River congeners by size of eye (Figs 11-15; Table 1), snout length,

and colour pattern. Its eye is easily recognisable as the largest.

Colour pattern features in which it differs from S. malleti are listed

under S. malleti. Distinguishing features for all described

Sundasalanx are presented in Table 4.

ECOLOGY. Sundasalanx megalops was captured at the mouth of a

creek over sandy, silty substrate in flowing turbid water approxi-

mately 1.5 m deep. Seining sand and mud bars along Sungai Joloi

up- and downstream from its point of capture produced specimens of

S. mesops.

DISTRIBUTION. Sundasalanx megalops was captured from a single

locality on Sungai Joloi, a large upper basin tributary of the Barito

River, Central Kalimantan, Indonesia.

ETYMOLOGY. This species is named megalops, a combination of

mega, Greek for large, and ops, Greek for eye.

Sundasalanx platyrhynchus Siebert and Crimmen, sp. nov.

(Fig. 15, 16)

HOLOTYPE. MZB 5944 (ex CMK 6979), 19.2 mm SL, Indonesia,

Kalimantan Barat, Kapuas River basin, Kapuas R. mainstream

Table 4 Summary table of diagnostic differences among described species of Sundasalanx.

D.J. SIEBERT

S. praecox S. microps S. platyrhynchus S. malleti S. mesops S. megalops

Ipb 3 tooth plate large small; 1—2 teeth small; 2—3 teeth large; row of teeth medium; teeth small; 4-5 teeth

along post. edge along post. edge

Ipb 4 tooth plate large small large large large large

Palatal and Bb | teeth _ present absent absent absent absent absent

Gill rakers on Ist arch well devel.; 8+1; minute; 2—3+0 minute; 2—3+0 minute; 5+1 minute; 3+0 minute; 2+0

along whole arch

Upper jaw teeth 10-11 + 30 5-9 + 15-19 5-7 + 15-19 4-5 + 15-17 3-5 + 16-18 2-3 + 12-14

Lower jaw teeth 16; 2 rows 15-16; 2 rows 15-18; 2 row post. 10-12 12-13 8-10; 2 rows post

'Vertebrae 37-38 41-42 41-42-43 40-42-44 40-4] 4]

'Branched dorsal rays 9-10 9-10 9-10-12 8-10-11 8-9-10 10

"Branched anal rays 13-14-15 15-16-17 14-16-17 13-14-17 13-14-16 14-15

Eye size medium; 4-5% SL _ small; appox.3% SL medium; 4-6% SL small; 3-4.5% SL medium; 4-6% SL large; approx. 6% SL

Prepectoral spot absent present present present present present

Cardiac chamber absent present present absent present present

pigmentation

Mid-ventral line absent present present present present present

Basipterygium absent absent absent present absent absent

pigment ring

' Median value italicised.

q re] ' ° '

about 7 km SW of Nanga Silat, approx. 0° 19'N 111° 45'E, 29-IV- DISCUSSION

1990, M.Kottelat et al.

PARATYPES. CMK 6979 (12), collection data as for holotype.

CMK 7898 (3), c&s, data as for holotype. CMK 6892 (8), Indonesia,

Kalimantan Barat, Kapuas River basin, Kapuas R. mainstream at

Teluk Ujung Bayur, approx. 0° 50'N 112° 45'E., 27-IV-1990,

M. Kottelat et al.

DIAGNOSIS. A photograph of S. platyrhynchus is presented as Fig.

15; selected morphometrics are summarised in Table | and vertebral

and fin-ray counts are summarised in Table 2 and Table 3. It is a

species of Sundasalanx with a large head (HL 19.6—23.3% SL, HW

10.2-12.0% SL), relatively large eye (4.2-5.7% SL, 21.7—25.9%

HL), and short snout (15.3—21.4% HL). Prepectoral mark present;

posterior wall of cardiac compartment marked with melanophores;

basipterygium without ring of melanophores. Vertebral number 41—

43 (median=43);dorsal-fin rays 9-12 (median=10); anal-fin rays

14-17 (median=16). Premaxilla with 5—7 teeth; maxilla with 15—19

teeth; dentary with 15—18 teeth, arranged in 2 rows in posterior part

of dental arcade; 3rd pharyngobranchial tooth plate small, with only

2 or 3 teeth; 4th pharyngobranchial tooth plate small, about 1/3 size

of pharyngobranchial 4, with principal row of larger teeth along

posterior edge.

Sundasalanx platyrhynchus is easily distinguished from its Kapuas

River congener by size of eye and pharyngeal dentition. The eye of

S. microps is much smaller (Table 4) and pharyngeal dentition in S.

microps is much reduced. Sundasalanx platyrhynchus can be distin-

guished from its Barito River congeners by size of eye and vertebral,

dorsal- and anal-fin ray counts. S. malleti has a smaller eye; S.

megalops has a larger eye. S. mesops has fewer vertebrae and dorsal-

and anal-fin rays.

DISTRIBUTION. Sundasalanx platyrhynchus is known from the

Kapuas River basin, West Kalimantan, Indonesia.

ETYMOLOGY. The name platyrhynchus is a combination of platys,

Greek for broad, and rhynchos, Greek for snout.

Sundasalanx is very small and much of its anatomy is suggestive of

the larvae of lower teleosts generally (as realised by Roberts, 1984).

Evolution of a tiny adult size among clupeiforms is not unusual; it is

known both in engraulids (Cervigon, 1982; Nelson, 1986; Roberts,

1981; Whitehead er al., 1988) and other clupeids (Poll, 1948;

Roberts, 1972; Whitehead, 1988; Whitehead & Teugels, 1985).

Among these a good anatomical description is available only for

Sierrathrissa leonensis (Whitehead and Teugels, 1985). However

‘youthful’ an impression small size lends to Sierrathrissa leonensis,

it nevertheless is more or less a miniature adult. The degree of

ossification, development of the intestinal tract, development of the

air bladder, fin positions, etc. are all indicative of at least a juvenile

stage in the life cycle, if not of an adult.

Sundasalanx on the other hand is both tiny and underdeveloped.

Its level of skeletal and somatic development (straight gut, small air

bladder, myotomes not extending ventrally to cover ventral portion

of body cavity, presence of ventral fin-fold, undifferentiated

cartilaginous plate supporting a rayless pectoral fin, etc.) is compa-

rable to that of other unmetamorphosed clupeid larvae. Migration of

the dorsal fin forward to a position characteristic of adults is

considered to mark the transition between larvae and adults in

clupeiforms. The positions of the dorsal fin, anus, and anal fin, in

absolute terms and relative to each other, of the largest Sundasalanx

specimens studied are similar to the positions reported for other

unmetamorphosed spratelloidins (Leis & Trnski, 1989) and there is

no difference in the positions of any of these structures between the

smallest and largest specimen for any species studied. Sundasalanx

appears to have attained the ability to reproduce at a stage of

development equivalent to larvae of other clupeids. If this assess-

ment of developmental stage of Sundasalanx is correct the size of

Sundasalanx may be unusual. Transition, or metamorphosis, in

other spratelloidins often begins at sizes of less than half that

attained by Sundasalanx.

Small size, lack of somatic development beyond that of the

RELATIONSHIPS OF SUNDASALANX

larvae of other clupeoids, and obvious acceleration of gonadal

maturation suggest Sundasalanx are progenetic. Whitehead &

Teugels (1985) attributed the small size of Sierrathrissa leonensis

to progenesis, but Sundasalanx exhibits a more extreme condition

of the syndrome than Sierrathrissa leonensis since S. leonensis is,

morphologically, a miniature ‘adult’. Among fishes perhaps only

Schindleria (Johnson & Brothers, 1993) matches Sundasalanx for

degree of progenesis. Both Sundasalanx and Schindleria

(Schindler, 1932; Gosline, 1959) are obvious larval forms that

have attained reproductive capabilities. This may be what sets

them apart from other small progenetic fishes, like Sierrathrissa,

with paedomorphic reductive tendencies.

Sundasalanx is of such small size an individual probably only

goes through one sexual cycle during its lifetime. Mature or matur-

ing specimens have been taken from the Barito River during February

and August, suggesting Sundasalanx might breed year around, a

reproductive phenomenon observed for other species of tropical

Bornean freshwater fishes (Roberts, 1989). Alternatively, it may

follow a bimodal breeding pattern tuned to rainfall patterns of the

region since February and August are months which correspond to

the two periods of low rainfall and low water levels of the annual

climatic cycle of Central Kalimantan. Larvae of marine clupeoids

can attain the known size of Sundasalanx within two months or less

if water temperatures are high enough and resources adequate

(Blaxter & Hunter, 1982). Conditions permitting, Sundasalanx prob-

ably is capable of completing its life cycle within a single wet—dry

cycle only half a year long, and thus might be capable of completing

two generations per annum.

Weitzman & Vari (1988) and Kottelat & Vidthayanon (1993)

observed that many miniature freshwater fishes are found in still or

slow-flowing waters. Within the Barito River basin Sundasalanx

was captured adjacent to strong currents and its apparent ability to

move in-shore at dusk within a riverine environment suggests it is an

exception to this generalisation, as is also one of the other miniature

clupeiforms (Thrattidion noctivagus Roberts, 1972; Whitehead,

1988). Barito River Sundasalanx also appears to be found only in

larger habitats. It was never taken in small tributary streams, many of

which were sampled.

ACKNOWLEDGEMENTS. The Indonesian Institute of Sciences (LIPI) is

gratefully acknowledged for permission to conduct research in Indonesia; Dr.

Soetikno Wirjoatmodjo, Dr. Dedy Darnedi, Dr. Arie Budiman, and Mr. Pak

Amir, all of the Research and Development Centre for Biology, Bogor, Java,

have supported research efforts in the Barito basin and their support is

gratefully acknowledged; The Worshipful Company of Fishmongers, the

Royal Society, and the Godman Fund are acknowledged for financial sup-

port; Project Barito Ulu is acknowledged for logistic support; Dr. Maurice

Kottelat kindly made Kapuas River materials available from his private

collection. Oliver Crimmen is thanked for his efforts in describing the new

species. Colin Patterson, Dave Johnson, Peter Forey, and Lance Grande all

made valuable suggestions toward improving the manuscript. Finally my

colleague and friend Agus Tjakrawidjaja is warmly acknowledged for his

personal attention to research on freshwater fishes in Indonesia.

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Issued 26 June 1997

Redescription of and lectotype designation for

Balistes macrolepis Boulenger, 1887, a senior

synonym of Canthidermis longirostris

Tortonese, 1954 and C. villosus Fedoryako,

1979 (Teleostei, Tetraodontiformes, Balistidae)

ANTHONY C. GILL

Division of Lower Vertebrates, Department of Zoology, The Natural History Museum, Cromwell Road, London

SW7 SBD, United Kingdom

JOHN E. RANDALL

Ichthyology Division, Bishop Museum, P.O. Box 19000A, Honolulu, Hawaii 96817-0916, U.S.A.

Synopsis. Balistes macrolepis Boulenger, 1887 is identified as a valid species of Canthidermis from an examination of the two

stuffed syntype specimens, 444 and 457 mm SL, from Muscat, Oman. The larger of these is designated lectotype. The species is

redescribed on the basis of the lectotype, paralectotype and two juvenile (30.5—32.0 mm SL) specimens from the Gulf of Aden,

and is shown to be a senior synonym of C. /ongirostris Tortonese, 1954 (described from a 365 mm SL specimen from the Dahlak

Islands, southern Red Sea) and C. villosus Fedoryako, 1979 (described from eight 55.5—177.3 mm SL juveniles from the Gulf of

Aden). Canthidermis macrolepis is distinguished from congeners in having fewer body scale rows (35-40 versus 39-58).

Juveniles of C. macrolepis are readily distinguished from those of congeners by scale morphology (relatively long, branched,

fleshy outgrowths present on body and head scales versus fleshy outgrowths short and unbranched or absent) and coloration (pale

spots on head and body large and forming a network pattern versus pale spots absent or small and not forming a network pattern).

INTRODUCTION

Boulenger (1887) described Balistes macrolepis from two large

(stated total length one foot Il inches), dry specimens from Muscat,

Oman. With the exception of two books [Randall (1995) and Debelius

(1996)] that give accounts for B. macrolepis resulting from the

present research, the species has not been referred to subsequently.

Boulenger gave the following characters for the species: dorsal-fin

rays III + 26; anal-fin rays 23; a groove in front of eye; no enlarged

scales behind the gill opening; no spines on the tail; falciform dorsal

and anal fins; and a strongly notched caudal fin. The absence of

enlarged scales behind the gill opening is unique among Indo-

Pacific balistids to the genera Canthidermis Swainson, Xanthichthys

Kaup and Xenobalistes Matsuura (Matsuura, 1980, 1981; Smith &

Heemstra, 1986). There are two stuffed specimens labelled as B.

macrolepis in the Natural History Museum, London, which we

believe to be the syntypes. One (BMNH 1887.11.11.334; Fig. 1)

measures 457 mm SL and is mounted on an exhibition stand,

whereas the other (BMNH 1887.11.11.335) measures 444 mm SL.

The specimens have a terminal mouth with uneven, notched teeth, a

deep groove before the eye, and a relatively well-developed third

dorsal spine extending above the dorsal edge of the body, and lack

enlarged scales behind the gill opening and longitudinal grooves on

the cheek; this combination of characters is unique among balistids

to species of the genus Canthidermis (Matsuura, 1980).

Recent authors (e.g., Berry & Baldwin, 1966; Matsuura, 1980;

Smith & Heemstra, 1986) have recognised only two valid species of

Canthidermis, the cosmopolitan C. maculatus (Bloch) and the At-

lanticC. sufflamen (Mitchill). However, Fedoryako (1979) recognised

five species of Canthidermis in the most recent review of the genus:

C. maculatus, C. sufflamen, C. willughbeii (Lay & Bennett) (from

the eastern Pacific), C. rotundatus (Proce) (from the Indo-West

Pacific), and C. villosus, which Fedoryako described as a new

species.

Fedoryako (1979) described C. villosus from eight pelagic juve-

niles (55.5—177.3 mm SL) from the Gulf of Aden. He distinguished

it from juvenile congeners in having relatively long, branched,

fleshy outgrowths on body and head scales (versus fleshy outgrowths

short and unbranched or absent), large, pale spots on head and body

forming a network pattern (versus pale spots absent or not forming

a network pattern), and 36-40 (versus 39-57) transverse rows of

scales (= body scale rows). We located two additional juvenile

specimens of the species in the Natural History Museum, London,

(BMNH 1939.5.24.1849-1850) that had been surface dipnetted in

the Gulf of Aden by the 1933-34 John Murray Expedition (Station

25) on the 10th of October, 1933. These specimens had been

identified as Canthidermis sp. and briefly described in Norman’s

(1939) report of fishes of the 1933-34 John Murray Expedition.

Adult specimens of C. villosus have not been described. However,

the fin-ray and scale counts of the two adult syntypes of Balistes

macrolepis agree closely with those of C. villosus (Table 1), and we

conclude that the two nominal species are conspecific. Balistes

macrolepis Boulenger, 1887 is therefore a senior synonym of

Canthidermis villosus Fedoryako, 1979.

The second author searched his photographic library and found

four photographs of individuals that we believe to be conspecific

with the syntypes of B. macrolepis. One photograph taken at Fahl

Island off Muscat in the Gulf of Oman by J.P. Hoover shows a

nesting pair (Fig. 2). Two others by H. Debelius are of specimens

from Oman, one a natural underwater photograph, the other of an

aquarium specimen in the Muscat Aquarium; both photographs

were reproduced in Debelius (1993: 298). The final photograph,

taken by the second author, is of aca. 300 mm total length specimen

in the Zubayr Islands, southern Red Sea (reproduced in Randall,

1995: fig. 1108). The localities for these photographs and specimens

suggest that the species might be restricted to the northwestern

Indian Ocean and Red Sea. We searched literature on Canthidermis

from the area and found two additional references. Tortonese (1954)

described C. longirostris from the Dahlak Islands, southern Red Sea,

and Dor (1984) recorded C. maculatus from the Red Sea. However,

Dor’s record was based on Tortonese’s specimen of C. longirostris,

and followed Berry & Baldwin’s (1966) synonymy of the two

species; apparently, Fedoryako (1979) overlooked C. longirostris in

his review of the genus. Examination of Tortonese’s description and

figure of C. longirostris revealed that it is not referable to C.

maculatus, rather it is a second junior synonym of B. macrolepis.

MATERIALS AND METHODS

Pectoral-fin ray counts include the uppermost, rudimentary ray.

Other methods of counting and measuring follow Matsuura (1980).

Institutional codes follow Leviton et al. (1985).

SYSTEMATICACCOUNT

Canthidermis macrolepis (Boulenger, 1887)

Figures 1—3; Tables 1—2

Balistes macrolepis Boulenger, 1887: 666 (type locality: Muscat;

lectotype: BMNH 1887.11.11.334, 457 mm SL, designated be-

low).

Canthidermis sp. — Norman, 1939: 109 (Gulf of Aden).

H 1

a eHeHeEHEEHEesl

A.C. GILL AND J.E. RANDALL

Canthidermis longirostris Tortonese, 1954: 77, fig. 1 (type locality:

Dahlak Is, Red Sea; holotype: MZGZ 20162, 365 mm SL).

Canthidermis villosus Fedoryako, 1979: 985, fig. 1B (type locality:

12°29'N 44°23’E, Gulf of Aden; holotype: MGY P-15097, 55.5

mm SL). — Fedoryako, 1981: 21, fig. 1c (English translation of

original description).

Canthidermis maculatus [non Balistes maculatus, Bloch 1786]. —

Berry & Baldwin, 1966: 463 (synonymy with C. longirostris). —

Dor, 1984: 275 (Red Sea; based on holotype of C. longirostris). —

Debelius, 1993: 298 (colour photographs of specimens from

Oman).

Canthidermis macrolepis. — Randall, 1995: 393, fig. 1108 (descrip-

tion, synonymy and distribution based on present study; colour

photograph). — Debelius, 1996: 298 (colour photographs of speci-

mens from Oman).

DIAGNOSIS. Canthidermis macrolepis is readily distinguished from

congeners in having fewer body scale rows (35-40 versus 39-58).

As noted by Fedoryako (1979, 1981; see above), juveniles of

Canthidermis macrolepis are readily distinguished from those of

congeners by scale morphology (relatively long, branched, fleshy

outgrowths present on body and head scales versus fleshy outgrowths

short and unbranched or absent; Fedoryako, loc. cit.: fig. 1C cf. fig.

1A,B) and coloration (pale spots on head and body large and

forming a network pattern versus pale spots absent or small and not

forming a network pattern).

DESCRIPTION. (based on data from BMNH specimens; see Tables

1,2 for counts and measurements of individual specimens, and for

data from Tortonese’s and Fedoryako’s respective descriptions of C.

longirostris and C. villosus)

Dorsal-fin rays If] + 25-26, all segmented rays branched except

for the first 1-2; anal-fin rays 22—23, all rays branched except for the

first; pectoral-fin rays 15-16, the upper ray a rudiment, and the

second from uppermost unbranched; body scale rows 35-39; head

scale rows 25—29; vertebrae 7 + 11 (from radiographs of BMNH

1939.5.24.1849-1850 only); gill rakers 8-10 + 20-22 = 30, the

Fig. 1 Canthidermis macrolepis, BMNH 1887.11.11.334, lectotype, 457 mm SL, Muscat, Oman (photograph by P. Hurst).

BALISTES MACROLEPSIS — REDESCRIPTION AND LECTOTYPE DESIGNATION 29

Fig. 3 Canthidermis macrolepis, BMNH 1939.5.24.1849—1850, 30.5 mm SL, 1933-34 John Murray Expedition station 25, Gulf of Aden (drawing by

A.C. Gill).

upper-lobe rakers markedly smaller than those of lower lobe (from

examination of right sides of BMNH 1939.5.24.1849-1850).

Body relatively deep in juveniles (greatest body depth 56.3-

57.0% SL; depth of body 46.6—-46.9% SL), more elongate in adults

(greatest body depth 32.9-35.9% SL; depth of body 27.9-29.3%

SL); width of body 22.3-22.5% SL in juveniles, 14.9-15.3% SL in

adults; head large in juveniles (head length 39.0-39.7% SL), shorter

in adults (26.0-27.5% SL); snout length 19.0-19.1% SL in juve-

niles, 17.1-17.8% SL in adults, the dorsal profile of snout slightly

concave in juveniles and slightly convex in adults; eye round, the

greatest orbit diameter 11.8—12.1% SL in juveniles, 4.6-4.7% SL in

adults; interorbital space broad (interorbital width 13.1% SL) and

concave in juveniles, convex and narrower (interorbital width 9.4—

9.5% SL) in adults; caudal peduncle short and deep in juveniles

(least depth 12.5-12.8% SL, length 12.8-13.1% SL), long and

slender in adults (least depth 8.3% SL, length 22.8% SL); pelvic flap

not capable of large ventral expansion, the free pelvic terminus with

two apparent, weakly movable sections.

Mouth small and terminal; teeth incisiform, sharp and notched on

the edges, projecting and close set, four on each side of jaws, with an

Table 1

A.C. GILL AND J.E. RANDALL

Meristic frequencies for specimens of Canthidermis macrolepis. Data for type specimens of Canthidermis longirostris and C. villosus follow

Tortonese (1954) and Fedoryako (1979), respectively.

D, rays A rays P, rays* Body scale rows* Head scale rows*

24 25 26 22 28. 2A sy iN 35 36) 3788, 23905 40) 25) 26 29

B. macrolepis

Lectotype - - 1 - 1 - ~ 2 - - - - 2 - 2 - -

Paralectotype - 1 - damaged 1 1 - p) - ~ - - 2 -

C. longirostris holotype - - i ~ - 1 not given - - - 1 - not given

C. villosus types** 1 D, 3 3 5 = 1 7 = - 3 1 D not given

Murray specimens - 2 - 1 i - 4 1 i 2 - - - - - 1

Totals 1 5) 5 4 7 1 2 14 l 3 4 4 3 2 2. 2

* - characters where bilateral counts are included for some specimens. ** - it is apparent from data given for other Canthidermis species that Fedoryako (1979) did not include

the upper rudiment in his counts of pectoral-fin rays; we have therefore added one to the values recorded by him.

Table 2 Selected morphometric values expressed as percentages of SL for specimens of Canthidermis macrolepis.

Murray specs Holo. Paratypes Paratypes longirostris macrolepis types

(n=3) (n=4) holo. Paral. Lecto.

SE 30.5 32.0 55:5 75-95 111-177 365 444 457

Head length 39.0 39.7 37.6 35.0-36.4 33.6-34.0 26.5 27S 26.0

Snout length 19.0 19.1 19.8 19.8-20.4 19.2—20.6 3 17.8 17.1

Body depth 46.6 46.9 Sa vs nah ae 278 298

Greatest body depth 57.0 56.3 53.0 46.2-53.3 41.1-48.3 ae 32.9 35.9

Body width 228) 22.5 oe Be ae Ss 14.9 1533

Snout to D, origin 43.6 43.8 42.0 40.3-42.3 35.2-38.9 ot 29.7 28.4

Snout to D, origin 65.9 63.4 a ee ie ot 54.5 54.5

Interdorsal space 23.6 20.6 ae oe css =e 26.4 27.6

Snout to A origin Syl iss 73.0 69.5-73.7 66.3-68.1 she 61.0 60.8

D, base length 29.5 30.3 28.8 29.4-29.8 27.9-30.2 ie 27.0 26.9

A base length 26.9 2S 24.2 24.3-24.7 23.9-24.9 ss ss 22.3

Gill opening length WE) 9.4 ae ae aly 6.0 4.7 4.8

Eye diameter 1), 11.8 10.5 8.4-8.8 7.1-8.9 4.9 4.7 4.6

Caudal peduncle length* 12.8 13.1 12.0 12.6-14.1 13.4-17.2 a cas 22.8

Caudal peduncle depth V3) 12.8 12.1 10.9-12.7 10.3-11.4 oD 8.3 8.3

First dorsal spine length 223 21.8 ae a Ae ve 11.0 us

Longest D, ray length 20.0 ANS) 24.0 20.2-24.8 20.7-29.5 at ve 22.5

Longest A ray length 18.7 20.3 20.4 20.5—22.7 22.0-25.4 nae ants 21.4

Pectoral fin length 15.4 16.3 £5 ace a 9.6 10.1 a

Caudal fin length PIAS) 23a sp ae ma 20.4 24.3 sah

C. villosus types

* - caudal peduncle length values given for the C. villosus types are the postdorsal distance values given by Fedoryako (1981; measured from base of last second-dorsal ray base

to base of caudal fin). ** - character not given in literature or not available because of specimen damage.

inner three plate-like teeth on each side of upper jaw. Gill opening

slightly oblique (sloping in posterodorsal direction), its length 7.9—

9.4% SL in juveniles, 4.7-4.8% SL in adults; no patch of modified

scales posterior to gill opening. Nostrils small, located just in front

of orbital rim between about 9 and 10 o’clock position from centre

of eye. A deep groove extending beneath nostrils from midanterior

edge of eye along upper third of snout. No longitudinal or diagonal

grooves on cheek.

Scales of juveniles rhomboidal, not overlapping, with a large

spine projecting posterolaterally from scale centre and well-devel-

oped ridges that extend posteroventrally and posterodorsally from

base of central spine; ridges usually ending dorsally and ventrally

with a smaller spine; scale spines each with a fleshy outgrowth, these

small and unbranched on most scales, but large and highly branched

on at least some scales. Scales of adults rhomboidal, weakly over-

lapping, those on body with a spine-like ridge on centre, and about

25-60 small nodules arranged in a diamond-shaped patch immedi-

ately behind ridge. Lateral line not apparent.

Origin of spinous dorsal fin about 1 (juveniles) to 2 (adults) eye

diameters posterior to eye; first dorsal-fin spine of juveniles stout, its

length 21.8—22.3% SL, with two ridges on its lateral surfaces, these

converging near base and tip of fin spine and bearing large, irregular

spines; first dorsal-fin spine of adults stout, its length 11.0% SL,

with nodules on its anterior surface, these small and arranged in

about 12 irregular rows proximally, becoming larger and arranged in

3 prominent rows distally; second spine of juveniles and adults

slender, without spinules or nodules, about half to two-thirds length

of first spine, acting as trigger to release the first spine when the latter

is locked in upright position; third spine of juveniles and adults

slender and short, about two-thirds length of second spine, without

spinules or nodules, and partly concealed by a deep groove into

which the spinous dorsal fin folds; origin of soft dorsal fin about one

eye diameter anterior to anal-fin origin in juveniles, and above anus

in adults; first soft dorsal-fin ray short, the longest ray in juveniles

the sixth or seventh, 20.0—21.3% SL, and in adults the fourth, 22.5%

SL; first anal-fin ray short, the longest ray in juveniles the fifth or

sixth, 18.7—20.3% SL, and in adults the fourth, 21.4% SL; soft

dorsal and anal fins broadly rounded in juveniles, strongly pointed

BALISTES MACROLEPSIS — REDESCRIPTION AND LECTOTYPE DESIGNATION 31

and falcate in adults; pectoral fins rounded, 15.4-16.3% SL in

juveniles, 10.1% SL in adults; caudal fin rounded in juveniles,

double emarginate in adults, its length 23.1—24.3% SL; segmented

fin rays of juveniles bearing 1—3 rows of small spinules, these largest

on basal part of fins; segmented-fin-ray spinules not apparent in

adults.

PRESERVED COLORATION OF JUVENILES (based on BMNH

1939.5.24.1849-1850; Fig. 3). Head and body pale brown, paler

ventrally; body and caudal peduncle with large (slightly larger than

pupil to about twice diameter of eye), pale brown to whitish spots

arranged in about six oblique columns; about six spots in anteriormost

column (just behind pectoral-fin base), reducing to two spots in

posteriormost column (on caudal peduncle); pale spots most con-

spicuous on caudal peduncle and posterior part of body; interspaces

between spots accentuated with dark grey-brown stripes, bars and

(particularly at junctions of bars and stripes) spots, these forming a

reticulate pattern; dark grey-brown markings most conspicuous

below first dorsal fin, on caudal peduncle and adjacent to bases of

anal and second dorsal fins; head with pale spots and irregular

markings, these generally smaller and less distinct than those on

body; fleshy scale outgrowths mostly unpigmented, except for a few

scattered dark grey-brown unbranched outgrowths; first dorsal fin

pale brown to hyaline, with three large dark grey spots, one behind

each fin spine; second dorsal and anal fins pale brown to hyaline,

with large pale spots and dark grey reticulate pattern extending on to

basal two thirds of fin; caudal fin pale brown to hyaline, dusky

basally near dorsal and ventral edges of fin, with a dusky bar through

proximal third of fin; pectoral fins pale brown to hyaline.

Fedoryako (1981: 22) gave the following description for pre-

served juveniles: ‘The coloration of preserved specimens up to 100

mm [SL] in size is beige with a coarse reticular pattern. The 2nd

dorsal, the anal and caudal fins, have large light spots along the

margin and the basal part. In large fish [i.e., 111-177 mm SL] the

head and trunk are of a uniform beige, the ventral surface of the head

and trunk in front of the pelvic fins is lighter.’

LIVE COLORATION OF ADULTS (see Debelius, 1993, 1996; Randall,

1995). Head and body grey, shading to pale ventrally; dorsal, anal

and caudal fins grey, the distal edges of caudal, anal and second

dorsal fins dark grey; pectoral fins dark grey.

DISCUSSION

We herein designate the larger syntype of Balistes macrolepis

(BMNH 1887.11.11.334, 457 mm SL) lectotype of the species.

Species of Canthidermis are pelagic, only coming into shallow

areas to breed, whereupon demersal eggs are laid in a large pit in

sand and/or rubble (Fig. 2). Juveniles are usually found in associa-

tion with floating debris and vegetation; Fedoryako (1979: 986;

1981: 21) indicated that the type specimens of C. villosus were

collected near the surface beneath ‘floating microphytes.’ Presum-

ably, juveniles are camouflaged by their spotted/reticulate colora-

tion and branched, fleshy scale outgrowths. Current evidence suggests

that C. macrolepis breeds in Oman from at least July to November.

J.P. Hoover has photographed adults of the species in shallow areas

in Oman during the months of July and September, and J.K.L. Mee

(pers. comm.) informed us that adults are commonly caught by

hook-and-line fishermen working in shallow areas off Muscat around

October to November. Interestingly, although taste tests conducted

by Omani fisheries researchers indicate that the species is one of the

most flavoursome of Omani fishes, Omani fisherman usually dis-

card their catches believing the species to be poisonous, or at least

inedible (J.K.L. Mee, pers. comm.).

ACKNOWLEDGEMENTS. Weare grateful to H. Debelius and J.P. Hoover for

making photographs of Canthidermis macrolepis available to us, and to

J.K.L. Mee for helpful discussions. We thank P. Hurst of the Photographic

Unit of the Natural History Museum for the photograph of the lectotype

reproduced in Fig. 1. P.-C. Heemstra kindly loaned comparative materials of

Canthidermis from various western Indian Ocean localities. We thank A.-M.

Woolger and S. Davidson for preparing radiographs of specimens, and O.A.

Crimmen for assistance in locating the syntypes of Balistes macrolepis. We

are grateful to J.B. Hutchins, K. Matsuura and D.J. Siebert for their construc-

tive reviews of the manuscript.

REFERENCES

Boulenger, G.A. 1887. An account of the fishes obtained by Surgeon-Major A.S.G.

Jayakar at Muscat, east coast of Arabia. Proceedings of the Zoological Society of

London 1887(43): 653-667, pl. 54.

Berry, F.H. & Baldwin, W.J. 1966. Triggerfishes (Balistidae) of the eastern Pacific.

Proceedings of the California Academy of Sciences (4)34(9): 429-474.

Debelius, H. 1993. Indian Ocean Tropical Fish Guide. 321 pp. Aquaprint Verlags

GmbH, Neu Isenburg.

—— 1996. Fischfiihrer Indischer Ozean. 321 pp. IKAN-Unterwasserarchiv, Frankfurt.

Fedoryako, B.I. 1979. Cnunoporu poxos Canthidermis u Xanthichthys (Balistidae,

Tedraodontiformes [sic]) THxoro HM MHAMACKOrO OKeaHOB. Voprosy [khtiologii

19(6): 983-995. [In Russian]

1981. Triggerfishes of the genera Canthidermis and Xanthichthys (Balistidae,

Tedraodontiformes [sic]) from the Pacific and Indian Oceans. Journal of Ichthyology

19(6): 19-30.

Leviton, A.E., Gibbs Jr, R.H., Heal, E. & Dawson, C.E. 1985. Standards in herpetology

and ichthyology: Part |. Standard symbolic codes for institutional resource collec-

tions in herpetology and ichthyology. Copeia 1985(3): 802-832.

Matsuura, K. 1980. A revision of Japanese balistoid fishes. 1. Family Balistidae.

Bulletin of the National Science Museum, Series A (Zoology) 6(1):27-69.

1981. Xenobalistes tumidipectoris, a new genus and species of triggerfish

(Tetraodontiformes, Balistidae) from the Marianas Islands. Bulletin of the National

Science Museum, Series A (Zoology) 7(4):191—200.

Norman, J.R. 1939. The John Murray Expedition 1933-34. Scientific Reports. Vol. 7,

no. |. Fishes. 116 pp. British Museum (Natural History), London.

Randall, J.E. 1995. Coastal Fishes of Oman. 439 pp. Crawford House Publishing Pty

Ltd, Bathurst.

Smith, M.M. & Heemstra, P.C. 1986. Family no. 263: Balistidae. pp. 876-882. In:

Smith, M.M. & Heemstra, P.C. (eds) Smiths’ Sea Fishes. Springer-Verlag, London.

Tortonese, E. 1954. Spedizone subacquea italiana nel Mar Rosso. Ricerche zoologische.

VI. Plettognati. Rivista di Biologia Coloniale 14: 73-86.

Bull. nat. Hist. Mus. Lond. (Zool.) 63(1):33-49

Issued 26 June 1997

A review of the Diogenes (Crustacea,

Paguridea) hermit crabs collected by Bedford

and Lanchester from Singapore, and from the

‘Skeat’ Expedition to the Malay Peninsula,

with a description of a new species and notes

on Diogenes intermedius De Man, 1892 GAN abu,

PATSY A. MCLAUGHLIN /,

Shannon Point Marine Center, 1900 Shannon Point Road, Anacortes, WA 98221-4042, U.S.A.

PAUL F. CLARK

Department of Zoology, Natural History Museum, Cromwell Road, London SW7 SBD, England.

CONTENTS

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SYNOPSIS. Species of the hermit crab genus Diogenes, collected by Francis P. Bedford and William F. Lanchester from

Singapore, and from the ‘Skeat Expedition’ to Malay Peninsula near the turn of the century have been reviewed for the first time.

Specimens identified as D. senex Heller from Lanchester’s Singapore collecting trip represent an undescribed species, while those

from the ‘Skeat’ expedition to the Malay Peninsula have proved to represent the taxon recently described as D. stenops Morgan

and Forest. Similarly, specimens identified by Lanchester as D. rectimanus Miers do not represent that taxon, but rather D. avarus

Heller and D. goniochirus Forest. Because of this latter confusion, the holotype of D. rectimanus has been redescribed.

Lanchester’s specimens assigned toD. planimanus Henderson include bothD. planimanus and D. intermedius De Man. However,

the species name, D. intermedius De Man, 1892 is preoccupied and a replacement name is proposed. Diogenes mixtus Lanchester

also has been redescribed, and D. desipiens Lanchester placed in synonymy with Paguristes hians Henderson. All of the species

have been illustrated.

INTRODUCTION

In a recent review of specimens assigned to Diogenes senex Heller,

1865, by McLaughlin and Haig (1995), Heller’s (1865) taxon was

restricted, by neotype designation, to a species presently known only

from the eastern coast of Australia.Among the other specimens of D.

senex sensu lato examined by McLaughlin and Haig (1995) were

three lots from the Natural History Museum (BMNH): one from

East Africa (BMNH 1955.3.5.58-60), one from the Suez Canal

(BMNH 1927.11.2.226), and one from Singapore (BMNH

1905.10.21.33-36). These authors did not attempt to identify the

species from East Africa. They noted that the specimen from the

Suez Canal agreed with those identified by Bouvier (1892) and

Nobili (1906) as D. senex from Suez and Djibouti that had subse-

quently been referred to D. gardineri Alcock, 1905 by Lewinsohn

(1969). While the specimens from Singapore ‘were definitely’ not

D. senex sensu stricto, McLaughlin and Haig were unable to deter-

mine their identity. All three lots have now been reexamined. The

specimens from the Suez Canal have been compared with specimens

of D. gardineri from its type locality, the Maldive Islands, and the

identity of the Museum specimens as that species has been verified.

The specimen from East Africa has also proved to be D. gardineri.

The specimens from Singapore represent an undescribed species.

There is little information accompanying these latter specimens

except the locality, Blakang mati, Singapore, and the collectors,

Bedford and Lanchester. However, as mentioned by Lanchester

(1900a) and Ingle (1991), Bedford and Lanchester collected in the

Straits Settlements (Singapore and Malacca) during a seven month

period (1899-1900). Francis P. Bedford was an echinoderm special-

ist, but collected on that occasion with Lanchester (Ingle, 1991).

William F. Lanchester is best known in the British carcinological

community for his publications on the Brachyura from Singapore

and Malacca, the Crustacea of Malaysia from the collections of the

Karawak Museum and those of the ‘Skeat’ Expedition to Malaysia

(Lanchester, 1900a, 1900b, 1901; Ingle, 1991). Lanchester’s (1900a)

report ‘On a collection of crustaceans made at Singapore and

Malacca. — Part I. Crustacea Brachyura’, suggested that a second

part, dealing with the other Crustacea, was planned, but apparently

was never published. Although the brachyuran crabs from the Singa-

pore collection were acquired by the Natural History Museum in

1900 (reg. 1900.10.22); these particular hermit crabs were not added

to the registry until 1905.

The second part of Lanchester’s report on the Crustacea from the

‘Skeat’ Expedition to Malaysia (Lanchester, 1902) did deal with the

Anomura, Cirripedia, and Isopoda, and included the descriptions of

two new species of Diogenes, i.e., D. desipiens Lanchester, 1902 and

D. mixtus Lanchester, 1902, and one lot each of specimens attributed

to Diogenes senex, D. rectimanus Miers, 1884, and D. planimanus

Henderson, 1893. McLaughlin and Haig (1995) expressed the belief

that Lanchester’s (1902) D. senex might correctly represent a spe-

cies of the Troglopagurus group of Diogenes (cf. Forest, 1952).

All of Lanchester’s (1902) Diogenes species from the ‘Skeat’

Expedition in the collections of the University Museum of Zoology,

Cambridge (UMZC) were examined during the present study and

despite a thorough search, the single male specimen upon which

Lanchester based his description of D. desipiens could not be

located. Correspondence between Lanchester and then Superin-

tendent S.F. Harmer, and a list of Crustacea from the expedition

identified by Lanchester appear in volume V of the University

Museum’s ‘History of the Collection’. Only D. desipiens is absent

from that list, which suggests strongly that it was never registered at

the Museum (R. Symonds, pers. comm.). Consequently, searches

were made of the collections of the BMNH and those of the National

University of Singapore. Both failed to locate Lanchester’s speci-

men, and it must be presumed that it is no longer extant.

As indicated above, the BMNH’s Singapore specimens attributed

to D. senex are described herein as a new species of Diogenes.

Lanchester’s (1902) Malay D. senex is, as suspected by McLaughlin

and Haig (1995), a species of the Troglopagurus group of Diogenes,

i.e., Diogenes stenops Morgan and Forest 1991. Lanchester’s (1902)

description of D. mixtus consisted of little more than a comparison

with three other species. It is now redescribed from the type materi-

als and one additional specimen from Malaysia. Both Lanchester’s

(1902) D. rectimanus and D. planimanus have been compared with

type material of their respective taxa. His D. rectimanus is repre-

sented by two different taxa, D. avarus Heller, 1865, and a species

subsequently described by Forest (1956) as D. goniochirus Forest;

his D. planimanus includes specimens of both D. planimanus and D.

intermedius De Man, 1892.

Notwithstanding the length of Miers’ (1884) description, neither

it nor his figures are particularly diagnostic, which may account for

Lanchester’s (1902) confusion. Consequently, the holotype of D.

rectimanus, also in the collection of the BMNH, is redescribed. De

Man’s (1892) original description of D. intermedius was based on

the comparison of a single specimen (sex not indicated) from Pare

Pare, Celebes (Sudawesi, Indonesia), that seemed to differ from the

P.A. MCLAUGHLIN AND P.F. CLARK

presumed type specimens of D. custos (Fabricius, 1798) and D.

miles (Fabricius, 1787), but incorrectly attributed to H. Milne

Edwards (1837). De Man (1892) was sufficiently uncertain about

the singularity of his specimen that he described it simply as

Diogenes sp. with the notation that should it prove to be a species

distinct from D. custos it should be called D. intermedius. Although

he provided a rather detailed description, it essentially indicated the

differences between his specimen and D. custos, and was not

accompanied by any illustrations. Until now, subsequent reports

have not been based on material. The nomenclatural status of the

species name Diogenes intermedius De Man, 1892, is compromised

because it is preoccupied by Diogenes pugilator var. intermedius

Bouvier, 1891. This matter is resolved by proposing a replacement

name for De Man’s taxon.

Although the type specimen of D. desipiens has not been located,

Lanchester’s (1902) description and illustrations have been care-

fully examined. It is our opinion that D. desipiens is not a species of

Diogenes, but rather of Paguristes, and in fact is a junior subjective

synonym of Paguristes hians Henderson, 1888. A comparison of

Henderson’s taxon, based on its holotype and supplemental mate-

rial, with Lanchester’s (1902) account, provides justification for our

synonymy.

MATERIALS AND METHODS

The new species of Diogenes comes from the Bedford and Lanchester

collection housed in the BMNH. The syntypes of Diogenes mixtus,

the Malaysian specimens assigned to D. senex, and the specimens

identified by Lanchester (1902) as D. rectimanus and D. planimanus

have been borrowed from the University of Cambridge collection.

The holotype of Diogenes rectimanus, the specimen herein desig-

nated lectotype of D. planimanus, the holotype of Paguristes hians,

two paralectotypes of D. goniochirus, and the additional specimen

of Diogenes mixtus from Kuching, Malaysia, are in the BMNH

collection. The holotype of D. intermedius has been borrowed from

the Instituut voor Taxonomische ZoGlogie (Zodlogisch Museum),

Universiteit van Amsterdam (ZMA). Four paratypes of D. stenops,

borrowed from the Western Australian Museum (WAM), have been

compared to verify our determination of Lanchester’s (1902) ‘D.

senex’. One measurement, shield length (SL), measured from the

midpoint of the anterior margin of the shield to the midpoint of the

posterior margin provides an indication of specimen size. Photo-

graphs all were taken with Nikon 35 mm cameras equipped with a

Medical Nikkor 1:5.6/F=200 or Micro Nikkor Auto 1:3.5/F=55 mm

lens.

SYSTEMATIC ACCOUNT

Diogenes inglei sp. nov.

(Figs la—e, 2a—d, 10a)

MATERIAL EXAMINED. Holotype. Ovigerous 9 (SL = 1.46

mm), in ‘shell’ which actually is fragment of worm tube; Blakang

mati, Singapore, 1899, collectors, F Bedford and W. F. Lanchester,

BMNH 1905.10.21.33. Paratypes. 29, 1 ovigerous 9 (SL = 1.13-

1.52 mm), same locality, date and collectors, BMNH

1905.10.21.34—36.

DIAGNOSIS. Shield surface (Fig. la) anteriorly and laterally

DIOGENES OF SINGAPORE AND THE MALAY PENINSULA

bce

Fig. 1 Diogenes inglei sp. nov., holotype ovigerous 2 SL = 1.46 mm,

BMNH 1905.10.21.33; a. shield and cephalic appendages; b. anterior

lobe of sternite of 3rd pereopods; c. Ist maxilliped; d. 6th abdominal

tergite, protopods of uropods and telson; e. telson.

weakly spinulose. Dorsal margins of branchiostegites with 3-6

small spines or spinules. Ocular peduncles swollen proximally and

narrowing distally to tapering corneae; overreached by antennular

peduncles. Ocular acicles with 1-3 strong and | or more smaller

spines. Intercalary rostriform process slender, reaching beyond

proximal half of acicle but not to tip of inner-most acicular spines; no

ventral spine. Antennal peduncles reaching to or slightly beyond

distal margins of corneae. Antennal acicle simple, with bifid termi-

nal spine and 2 widely-spaced spines on mesial margins. Antennal

acicles with pair of long pinnate setae on each article. First maxilliped

(Fig. 1c) without exopodal flagellum.

Left cheliped (Figs 2a, 10a) with row of prominent spines on

upper margins of dactyl, palm and carpus, and lower margins of

palm and fixed finger; outer surface of dactyl with short row of

spines near upper margin; palm and fixed finger with scattered small

spines or tubercles, irregular row of larger spines near upper margin

of palm. Right cheliped (Fig. 2b) with row of 3 spines adjacent to

upper margin of palm; 3 strong spines on upper margin of carpus.

Carpi of ambulatory legs (Figs 2c, d) each with dorsodistal spine and

1 additional spine on dorsal surface proximally on second pereopods.

Tergite of sixth abdominal somite with lateral and terminal spines

marginally.

Telson (Figs 1d, e) with median cleft practically obsolete; termi-

nal margin with row of several large and few smaller spines, extending

onto lateral margins.

Fig. 2 Diogenes inglei sp. nov., holotype ovigerous 9 SL = 1.46 mm,

BMNH 1905.10.21.33; a. left chela; b. right chela; c. right 2nd

pereopod; d. left 3rd pereopod.

DESCRIPTION. Shield (Fig. 1a) longer than broad; anterior margin

between rostrum and lateral projections weakly concave; anterola-

teral margins with marginal or submarginal row of small blunt or

acute spinules; anterolateral angle unarmed; posterior margin trun-

cate; dorsal surface with scattered spinules anteriorly and laterally.

Rostrum obsolete. Lateral projections obtusely triangular, with promi-

nent marginal or submarginal spine. Dorsal margin of branchiostegite

with 3-6 sometimes widely-spaced small spines or minute spinules,

] spine on anterior margin.

Ocular peduncles moderately long, approximately 0.80 length of

shield; swollen proximally and narrowing distally to somewhat

reduced and distally tapered corneae, dorsomesial surface with row

of long fine setae in proximal half. Ocular acicles almost

subtriangular, anterior margin with 1—3 strong and 1 or more smaller

spines; separated basally by width of intercalary process. Latter

moderately well developed, somewhat depressed, reaching beyond

proximal half of ocular acicles, with terminal spinule; no ventral

spine.

Antennular peduncles, when fully extended, overreaching cor-

neae by approximately 0.33-0.50 length of ultimate segment.

Ultimate segment with 1 or 2 long setae on dorsodistal margin.

Penultimate segment with few scattered setae. First segment fre-

quently with small spine on ventrodistal margin.

Antennal peduncles reaching to, or slightly overreaching distal

margin of corneae; with supernumerary segmentation. Fifth seg-

ment with 3 or 4 pairs of long setae on ventral margin distally. Fourth

and third segments with few short setae. Second segment with

Fig. 3 Shield and cephalic appendages, a. Diogenes goniochirus Forest,

1956, 9 SL = 2.15 mm, UMZC Nov. 30, 1899; b. Diogenes avarus

Heller, 1865,o° SL = 1.53 mm, UMZC Nov. 30, 1899.

dorsolateral distal angle weakly produced and terminating in small

spine, dorsomesial distal angle with small spine, laterodistal margin

usually with 2 small spines, and | or 2 prominent slender spines

ventrally. First segment with 1 small spine on lateral margin ventrally.

Antennal acicle not reaching to distal apex of fourth peduncular

segment, terminating in bifid spine, with 2 widely-spaced spines on

mesial margin. Antennal flagella moderately short, approximately

as long as ambulatory legs; each article with pair of long pinnate

(pappose) setae.

Maxillule with endopod lacking external lobe. First maxilliped

(Fig. 1c) lacking flagellum; endopod fused to exopod. Third

maxilliped with 2 strong spines on basis, ischium without crista

dentata but with 1 very strong curved spine; merus with | or 2 spines

on ventral margin.

Left cheliped (Figs 2a, 3a) with dactyl approximately 0.33 longer

than upper margin of palm; cutting edge with row of calcareous

teeth; terminating in small calcareous claw, overlapped by fixed

finger; outer surface flattened, with short row of 4 or 5 blunt to

extremely slender and acute spines near upper margin and 2 or 3

spinules centrally; upper margin with row of strong, subacute to

extremely acute spines, decreasing in size distally and sparsely

interspersed with long setae; inner surface with few widely scattered

long setae. Fixed finger with outer surface flattened, armed with few

scattered small blunt or sharp spines; lower margin armed with row

of strong, blunt or acute spines, sparsely interspersed with long

setae, and forming weak curve with lower, similarly armed margin

of palm; cutting edge with row of calcareous teeth; terminating in

prominent calcareous claw. Palm with outer surface convex, armed

with scattered blunt or acute spines, with irregular row of stronger

spines beginning near midpoint of proximal margin, curving up-

ward, and continuing to near distal articulation with fixed finger;

upper margin with 5 or 6 strong subacute or very acute curved

spines; inner surface with few scattered tufts of short setae. Carpus

P.A. MCLAUGHLIN AND P.F. CLARK

approximately equal to or very slightly longer than palm; upper

margin with row of 4 or 5 strong spines, outer face convex and with

slightly oblique row of 3 widely-spaced spines; inner surface gla-

brous. Merus triangular; dorsal margin with row of spinules and tufts

of setae, 1 much stronger spine at dorsodistal margin; ventromesial

with 3 or 4 small spines in proximal half; ventrolateral margin with

3 or 4 spinules in distal half. Ischium unarmed.

Right cheliped (Fig. 2b) appreciably smaller than left. Dactyl

approximately equal to length of palm; upper margin not distinctly

delimited, outer surface with few spinules partially obscured by long

setae; cutting edge with row of fine corneous teeth, terminating in

small corneous claw and overlapped by fixed finger. Palm with row

of 3 spines and long setae on or adjacent to upper margin, convex

outer face with varying amounts of long setae, usually | additional

small spine distally near upper margin and second small spine near

base of fixed finger, lower margin unarmed; fixed finger with row of

very fine spinules in midline; cutting edge with row of corneous

teeth, terminating in small corneous claw; inner surfaces of dacty]l,

fixed finger and palm with long setae, most numerous on dacty! and

fixed finger. Carpus with long setae and 3 strong spines on upper

margin, outer surface convex, with few long setae and 1 prominent

spine at mid-distal margin; inner and lower surfaces with scattered

setae. Merus triangular; dorsal margin with 1 or 2 spinules and long

setae, | more prominent spine at dorsodistal margin; ventromesial

margin with 2 or 3 small spines in distal half, ventrolateral margin

with 1—3 spinules distally. Ischium unarmed.

Ambulatory legs (Figs 2c, d) with dactyls approximately equaling

length of propodi, both dactyls and propodi of second appreciably

longer than third, unarmed but with numerous long setae. Carpi with

dorsodistal spine and | spine on dorsal surface in proximal half

(second), unarmed or with tiny proximal spinule (third), dorsal and

ventral surfaces with long setae. Meri with long setae on dorsal and

ventral surfaces, second with 2 widely-spaced spines, third unarmed.

Ischiaunarmed, but with long dorsal and ventral setae. Sternite of third

pereopods with subquadrate to subrectangular anterior lobe (Fig. 1b),

terminal margin with 3-8 small spines and long setae.

Male pleopods unknown. Female with pleopods 24 well devel-

oped, biramous; pleopod 5 appreciably reduced, with rudimentary

exopod. Tergite of sixth abdominal somite with deep transverse

median furrow; posterolateral margins each with 2 spines, terminal

margin with | spine one each side of midline. Protopods of uropods

(Fig. 1d) each with row of small tubercles. Telson (Figs 1d, e) with

median cleft obsolete or only faintly indicated; terminal margin

slightly concave, with row of several large and few smaller spines,

extending onto lateral margins.

COLouR. Unknown.

DISTRIBUTION. At present recognized only from Singapore.

ETYMOLOGY. _Itis with great pleasure that we dedicate this species

to Ray Ingle, formerly of the Crustacea Section, the Natural History

Museum, in recognition of his many contributions to our knowledge

of decapod crustaceans in general, and of the Paguridea of the

northeastern Atlantic and Mediterranean regions in particular.

AFFINITIES. Diogenes inglei sp. nov. most closely resembles D.

gardineri in the general shapes of the left and right chelipeds, and in

the armature of the pereopods. Diogenes inglei sp. nov. is readily

distinguished from Alcock’s species by its longer antennal pedun-

cles and flagellum which carries a pair of long pinnate setae ventrally

on each article. Differences are also apparent in the armature of the

chelipeds. However, variation in cheliped armature is common in

species of this genus, and our present knowledge of D. inglei sp. nov.

is too limited to permit evaluation.

DIOGENES OF SINGAPORE AND THE MALAY PENINSULA

REMARKS. The first maxilliped of Diogenes species is not often

described or illustrated, but in those few species for which it has

been (e.g., Tirmizi and Siddiqui, 1982a), a two-segmented exopodal

flagellum is typical. Diogenes inglei sp. nov. is the only species of

the genus currently known to lack the flagellum; in its place are a

pair of short setae (Fig. 1c). Whether this loss is unique to D. inglei

sp. nov. or simply reflects inadequate examinations of other species

remains to be determined. It should be noted, however, that the only

hermit crabs routinely lacking a flagellate first maxilliped are

parapagurids or coenobitids. Diogenes inglei sp. nov. also appears

distinctive in having the sixth abdominal segment armed with spines

on the posterolateral and posterior margins. This again may simply

reflect inadequate observations of other species.

Diogenes rectimanus Miers, 1884

(Fig. 10b)

Diogenes rectimanus Miers, 1884:262, pl. 27, fig. c; Gordan,

1956:318 (in part; lit.).

? Diogenes rectimanus: Henderson, 1893:419; Alcock, 1905b:71,

pl. 6, figs 8, 8a, pl. 7, fig. 2, 2a; Ajmal Khan and Natarajan,

1984:20, fig. 17; Morgan, 1987b:175; Haig and Ball, 1988:167;

Rahayu and Forest, 1995:395.

Non Diogenes rectimanus: Lanchester, 1902:366.

MATERIAL EXAMINED. Holotypec’(SL = 4.30 mm), Prince of

Wales Channel, Torres Strait, BMNH 1882.7.

REDESCRIPTION. Shield slightly longer than broad, almost

subquadrate; anterior margin between rostrum and lateral projec-

tions slightly concave, with 5 or 6 small tuberculate spinules near

bases of lateral projections; anterolateral margins sloping; posterior

margin truncate; dorsal surface with few transverse spinulose ridges

laterally. Rostrum broadly rounded; lateral projections each with

rather strong marginal spine. branchiostegial margins each with 5 or

6 moderately well developed spines.

Ocular peduncles approximately 0.80 length of shield, moder-

ately stout; cornea not dilated; ocular acicles with straight inner

margins, broadly rounded anterolaterally, with 3 small, but promi-

nent spines and several additional smaller spinules, not extending

entire length of terminal margin. Intercalary rostriform process

reaching approximately to distal third of ocular acicles, broad

basally, tapering to moderately slender subacute tip.

Antennular peduncles overreaching corneae by almost entire

length of ultimate segment. Ultimate segment with few setae dorsally

and tuft distally on both ultimate and penultimate segments; basal

segment unarmed.

Antennal peduncles overreaching distal margin of cornea by

0.20-0.35 length of ultimate segment. Fifth segment with row of

tufts of setae on ventral margin; fourth segment with few scattered

setae and small spine on dorsolateral distal margin; third segment

with tuft of setae; second segment with small spine at dorsolateral

distal angle and very small spine on dorsomesial distal angle,

ventrolateral distal angle with small spinule; first segment with row

of small spinules on distal margin laterally and similar row of small

spinules on lateral margin ventrally. Antennal acicle not reaching to

apex of fourth segment, with strong bifid spine and 4 accessory

strong spines on mesial margin, few tufts of setae on lateral margin.

Maxillule without external endopodal lobe.

Dactyl of left cheliped (Fig. 3b) approximately 0.35 longer than

palm; double row of spines on upper margin, outermost strongest,

row of equally strong spines adjacent to upper margin, outer surface

with scattered small spinose tubercles and very short setae; cutting

edge with multiple series of small calcareous teeth, largest in distal

37]

third; inner surface with row of low protuberances. Palm slightly

shorter than carpus; upper surface with irregular triple row of spines,

outer surface with slightly concave area just below upper margin

with few scattered spinules and tubercles and tufts of short setae,

upper outer face with adjacent longitudinal row of moderately strong

spines in proximal half, not reaching articulation of palm, remaining

outer surface with rather widely-spaced small spines; proximal

margin with row of stronger spines continuing to lower margin,

lower outer surface of palm spinose, to with short setae; inner surface

with few very small tubercles in lower half; lower margin with row of

strong, outwardly directed spines, decreasing in size on fixed finger,

and adjacent second row of much smaller spines; surface of fixed

finger with spinulose tubercles. Carpus with row of moderately blunt

spines on upper margin (distal 2 or 3 stronger), outer surface convex,

with series of small tuberculate spines, lower margin with strong

spine at lower distal angle, inner and lower surfaces with few

tubercles. Merus triangular; dorsal surface rounded, with transverse

rows of small spines or spinules continuing onto lateral surface

dorsally, one such row adjacent to laterodistal margin appreciably

longer, mesiodistal margin with row of tuberculate spines, mesial

face tuberculate in ventral half, ventromesial margin with row of

broad, low bifid spines, ventrolateral margin with row of spines

distally becoming spinulose tubercles proximally, ventral surface

with numerous low bifid spinules. Ischium with row of small tuber-

cles on laterodistal margin ventrally and proximal margin.

Right cheliped with dactyl approximately 0.35 longer than palm;

upper surface with double row of small spines, outer surface with

row of slightly larger spines, both surfaces generally concealed by

long setae; cutting edge with row of calcareous teeth, terminating in

calcareous claw, slightly overlapped by fixed finger. Palm approxi-

mately 0.65 length of carpus; upper surface somewhat spinulose,

small spines or spinules forming quasi transverse rows; outer face

with few small spines or low protuberances and tufts of setae; fixed

finger with 2 rows of moderately strong spines on outer surface,

cutting edge with quite prominent calcareous teeth; inner surface of

palm with tufts of setae, row of widely-spaced tubercles on fixed

finger. Carpus with row of spines on upper margin and second

adjacent row on upper outer face, outer distal margin with 2 spines,

low protuberances on lower outer face, surfaces all with long setae.

Merus with short transverse spinulose ridges and tufts of setae on

dorsal margin, lateral face with low spinulose protuberances; vent-

rolateral margin with 2 acute spines distally, low sometimes bifid

spinulose protuberances proximally extending onto ventral surface;

mesial face generally glabrous, ventromesial margin with row of

small spinules distally, double row of stronger spinules proximally.

Ischium with 2 spinules on ventromesial margin.

Ambulatory legs similar. Dactyls 0.25—0.35 longer than propodi;

dorsal margins of dactyls each with almost double row of long stiff

dense setae, lateral faces with faint transverse sulcus in proximal

half and row of long setae, ventral margins each with row of long

setae proceeding onto lateroventral margin distally, mesial faces

also with row of long stiff dense setae ventrally and second row of

stiff setae beginning in upper half and progressing ventrally toward

claw. Propodi approximately 0.25 longer than carpi, right with

dorsal, ventral, mesial and lateral tufts of setae, left with tufts of

setae accompanied by row of spinules dorsal surfaces, strongest on

third. Carpi each with row of acute spines, somewhat shorter on

third. Meri with dorsal and ventral tufts of setae; second with

spinulose protuberances ventrally on lateral faces and row of small

spinules on ventral margin, ventrolateral distal margin also with row

of spinules. Ischia with 2 or 3 spinules (second) or unarmed (third).

Sternite of third pereopods with indistinctly bilobed anterior lobe,

each pseudo-lobe with tuft of setae.

Telson with small median cleft, terminal margins of both lobes with

long spines interspersed with slightly smaller spines, extending

down lateral margin of left only.

CoLour. Not known.

DISTRIBUTION. Known with certainty only from the Torres Strait,

Arafura Sea; ? Persian Gulf, Gulf of Aden, India, Sri Lanka,

Indonesia, and Northern Australia.

REMARKS. Henderson’s (1893) very brief diagnosis of D.

rectimanus could refer to several species. The fact that he noted that

the lower margin of the left chela and fixed finger was straight, and

the fingers very short suggests that he may not have been dealing

with Miers’ (1884) species. Alcock (1905) stated in his diagnosis

that the merus of the left cheliped was not spinose. The merus of the

left cheliped of the holotype of D. rectimanus has a spinulose dorsal

margin; both the ventromesial and ventrolateral margins are spinose.

However, Alcock’s (1905, pl. 6, fig. 8, pl. 7, fig. 2) figures show a

very strongly armed merus, and in other respects do bear consider-

able similarity to the holotype of D. rectimanus. None of the other

citations of this species are sufficiently detailed to ascertain whether

or not the authors were actually dealing with Miers’ (1884) taxon.

However, as pointed out by Rahayu and Forest (1995), Haig and

Ball’s (1988) notation on the reduced armature of the lower margin

of the palm of the left cheliped, does suggest that they may not have

been, despite the close proximity of their specimen to the type

locality.

Diogenes goniochirus Forest, 1956

(Figs 3a, 8a, 9a, 11a)

2mm

Fig. 4 Shield and cephalic appendages, Diogenes planimanus

Henderson, 1893, 9 SL = 4.85 mm, UMZC Nov. 30, 1899.

P.A. MCLAUGHLIN AND P.F. CLARK

Imm

Fig.5 Anterior portion of shield and cephalic appendages, Diogenes

platvoeti nom. nov., holotype SL = 4.81 mm ZMA De201.872.

Diogenes rectimanus: Lanchester, 1902:366 (in part) [Non Diogenes

rectimanus Miers, 1884].

Diogenes goniochirus Forest, 1956:527, figs S—7; Rahayu and For-

est, 1995:395.

MATERIAL EXAMINED. Type material. Paralectotypeo’ (SL = 2.78

mm), ovigerous 9 (SL = 3.05 mm), Long Hai, Cochinchine, Viet-

nam, Modest, 1905; BMNH 1995.1663-64.

Lanchester material examined. 1 9 (SL = 2.15 mm), ‘Loc.

?’, ‘Skeat’ Expedition, Malay Peninsula; UMZC, Nov. 30, 1899.

DIAGNOSIS. Shield (Fig. 3a) as broad or slightly broader than long;

anterior margin with only few spinules between broadly rounded

rostrum and slightly produced lateral projections; dorsal surface

with few transverse, setose and/or spinulose ridges laterally. Dorsal

margin of branchiostegite with row of closely-spaced small spines

over entire length. Ocular peduncles 0.75—0.80 length of shield;

corneae dilated little if at all. Ocular acicles broadly sub-triangular;

terminal margins with 3 or 4 spines, extending approximately half

length of margins. Intercalary rostriform process subovate, acute,

not reaching to tips of acicular spines, no ventral spine. Antennular

and antennal peduncles approximately equal in length, both over-

reaching ocular peduncles. Antennal acicles not forked, with simple

terminal spine, lateral margin with 1 spine distally, mesial margin

with row of 4-6 spines. Antennal flagella with long ventral setae.

Left cheliped (Fig. 11a) with ventral and lateral faces of merus

spinulose, ventromesial distal margin with row of very small spines;

outer face of carpus spinulose, with longitudinal row of small spines

centrally, culminating in strong spinose protuberance distally, upper

margin with row of spines; lower surface and margin of palm and

fixed finger straight or convex, armed with 3 to several rows of blunt

or spinulose tubercles, outer surface of palm spinulose but without

median crest, row of small spines on upper margin of palm, some-

times more prominent distally, double row of spines on dactyl. Right

DIOGENES OF SINGAPORE AND THE MALAY PENINSULA

— _

—-

——

Dany:

ee aon

NWaacs

2mm

Fig.6 Shield and cephalic appendages, Diogenes platvoeti nom. nov.,

9 SL = 4.67 mm, UMZC Nov. 30, 1899.

cheliped with row of very small spines on upper surfaces of carpus,

palm, and dactyl, all practically obscured by rows of long setae.

Ambulatory legs with dorsal margins of carpi each with row of

closely-spaced small spines; dorsal margins of propodi each with

row of very small spinules and long setae (second) or double row of

long stiff setae (third); mesial faces of dactyls (Fig. 8a) each with 4

rows of setae, dorsal and ventral rows long and simple, median rows

shorter and pinnate, more distinct on third.

Telson with distinct median cleft, lobes slightly asymmetrical;

terminal margins with 2-4 moderate to strong and 3-6 smaller

spines, sometimes extending onto lateral margins.

COLOUR. Not reported.

DISTRIBUTION. Vietnam; Malaysia; Indonesia.

REMARKS. The Malay specimen of D. goniochirus is the largest of

12 specimens identified by Lanchester (1902) as Diogenes

rectimanus, and the only one he removed from its shell. Lanchester

remarked that the small size of the specimens probably accounted

for the lack of prominence of the spines on the lower margin of the

left chela and more obscure arrangement of granules on this append-

age. Judging from the development of the pleopods, this specimen is

most probably a mature female. In having a prominent spinose

protuberance on the carpus and lack of a crest on the outer surface of

the palm of the left cheliped, this specimen agrees well with the

paralectotypes of D. goniochirus that we examined. The spinose

dorsodistal angle of the palm is also apparent, but not as prominent

2mm am

Fig.7 Shield and cephalic appendages, a. Diogenes stenops Morgan and

Forest, 1991, ovigerous 9 SL = 3.11 mm, UMZC Nov. 30, 1899; b.

Diogenes mixtus Lanchester, 1902, lectotypeo’ SL = 6.88 mm, UMZC

1.10050.

as in the paralectotype female. The setation of the mesial faces of the

ambulatory legs is a little more distinct, but corresponds well with

that of the paralectotypes. The remaining 11 specimens, all of

appreciably smaller size but none the less mature, differ markedly

from this specimen of D. goniochirus, and are assigned toD. avarus.

Diogenes avarus Heller, 1865

(Figs 3b, 8b, 9b, c, d, 1 1b)

Diogenes avarus Heller, 1865:83, pl. 7, fig. 2; Alcock, 1905: 68, pl.

6, figs 6, 6a; Forest, 1956:524, figs 14; Lewinsohn, 1969:37, fig.

4, Tirmizi and Siddiqui, 1982a:54, fig. 29; Haig and Ball,

1988:167; Rahayu and Forest, 1995:398, Figs 2b, g, h.

Diogenes rectimanus: Lanchester, 1902:366 (in part) [Non Diogenes

rectimanus Miers, 1884].

? Diogenes avarus: Ajmal Khan and Natarajan, 1984:18, fig. 15.

MATERIAL EXAMINED. Lanchester’s material. 70°, 39, 1 ovi-

gerous 9 , (SL= 1.04—1.52 mm), ‘Loc. ?’, ‘Skeat’ Expedition,

Malay Peninsula, UMZC, Nov. 30, 1899.

DIAGNOSIS. Shield (Fig. 3b) longer than broad, with few short

transverse spinulose ridges and long setae on dorsal surface; rostrum

obsolete or broadly rounded. Dorsal margins of branchiostegites

with 5S—8 small spines. Ocular peduncles short and moderately stout;

overreached by both antennular and antennal peduncles. Ocular

acicles broad, with 1—3 strong spines and several minute spinules on

terminal margin, not extending entire length. Intercalary rostriform

process slender, reaching beyond proximal half of acicle but not

beyond tip of inner-most acicular spines.Antennal peduncles slightly

shorter to nearly equal length of antennular peduncles. Antennal

acicle not reaching to distal apex of fourth peduncular segment, with

simple or bifid terminal spine, lateral margins usually 1 or 2 spines

distally, mesial margins with 3—7 small spines. Antennal flagellum

with paired long setae ventrally.

Left cheliped (Fig. 11b) with 1 or more rows of small simple or

bifid spines on dorsal margin of merus, ventromesial distal angle

with 3 or 4 prominent spines, ventrolateral margin with 2—5 much

smaller spines distally; broad upper surface of carpus with 3 rows of

small acute or subacute spines, stronger on slightly produced distal

angle, outer face angularly convex with tuberculate or spinose

protuberance at median distal margin, surface armed with blunt or

spinulose tubercles and small spines; lower margin of fixed finger

and palm straight, with irregular rows of small tubercles or subacute

spines; palm with convex outer surface armed with moderately to

closely-spaced tubercles, subacute or acute spines or spinules, and

with crest of stronger tubercles or spines proximally near midpoint

of proximal margin but not continued to articulation with dactyl,

upper margin with irregular usually double row of small spines,

strongest on produced upper distal angle; upper surface of dactyl

with 3 rows of spines. Right cheliped with noticeable hiatus between

dactyl and fixed finger; upper margins of carpus, palm and dacty]

each with 1-3 rows of small spines partially obscured by long

plumose setae. Ambulatory legs with dorsal margins of carpi each

with double row of small spines on dorsal surface of second, usually

only single row of smaller spinules on third; propodi with irregular

row of small spines or spinules, always on second, frequently on

third; mesial faces of dactyls (Fig. 8b) each with 2 rows of rather

widely-spaced moderately short setae. Anterior lobe of sternite of

third pereopods (Fig. 9b) with roundly rectangular, with tuft of setae

on either side of midline.

Telson (Figs 9c,d) with median cleft; terminal margin of left lobe

with 3-6 large spines extending onto lateral margin and several very

small spinules medianly, right terminal margin with 4-6 small

spines.

COLOUR. Carapace rather uniform dark brown. Ocular peduncles

uniform light brown, or brown with broad, oblique white band

medially; cornea golden. Proximal segment of antennular peduncles

dark brown; distal segment with brown chromatophores on white.

antennal flagella banded brown and white. Merus and carpus of left

cheliped solid light brown, chela white; right cheliped solid light

brown. Meri of ambulatory legs white with 2 brown bands; carpi

light brown with white band distally; propodi white medially, other-

wise dark brown proximally and light brown distally; dactyl dark

brown proximally and white distally (after Haig and Ball, 1988).

DISTRIBUTION. Indian Ocean, from Red Sea and east coast of

Africa to Mergui Archipelago; Malaysia; Vietnam; Philippine Is-

lands; Indonesia; New Guinea; northeast coast of Australia.

REMARKS. Lanchester’s specimens agree well with Forest’s (1956)

description and with small specimens of this species from northern

Australia, particularly in having spinules on the dorsal surfaces of

both the carpi and propodi of the ambulatory legs. This is in contrast

to the figures of D. avarus given by Rahayu and Forest (1995: Fig.

2 g, h) in which the propodi are unarmed, and the third left pereopod

has only a few dorsodistal spines on the carpus. However, we

observed a similar lack of propodal armature and reduced carpal

spination in specimens from Barunda Beach, Lovina Bali. Rahayu

and Forest’s illustrated specimen was one of their largest males (SL

= 3.5 mm), while our specimens from Bali had SL’s of 2.84—2.96

mm. The largest of the Lanchester specimens had a SL of only 1.52

mm. It may be that propodal spination is lost withincreased animal

P.A. MCLAUGHLIN AND P.F. CLARK

size, as the specimens agree in other morphological characters. The

identification of D. avarus by Ajmal Khan and Natarajan (1894:18,

fig. 15) is uncertain.

Diogenes planimanus Henderson, 1893

(Figs 4a, 9c,e, 12a)

Diogenes planimanus Henderson, 1893:416, pl. 39, figs 5, 6;

Lanchester, 1902:365 (in part); Nobili, 1903a:15; Dechancé,

1964: 35; Tirmizi and Siddiqui, 1982a:43, figs 21, 22; 1982b:fig.

18.

Diogenes custos var. planimanus: Alcock, 1905b:66, pl. 6, fig. 3;

Sundara Raj, 1927:133; Kamalaveni, 1950:80; Gordan, 1956:317.

? Diogenes planimanus: Ajmal Khan and Natarajan, 1984:16, fig.

ihe

MATERIAL EXAMINED. Lectotype (herein designated). 9 (SL=5.7

mm) Madras; BMNH 1894:7:21:4. Type locality restricted by

lectotype designation to Madras.

Lanchester material examined. 3 oO, 29 (SL = 4.72-6.36 mm)

Patani; UMZC, Nov. 30, 1899.

DIAGNOSIS. Shield (Fig. 4) nearly as broad as long; anterior

Fig. 8 Dactyl of left 3rd pereopod (mesial view), a. Diogenes

goniochirus Forest, 1956, 9 SL = 2.15 mm, UMZC Nov. 30, 1899; b.

Diogenes avarus Heller, 1865, SL = 1.53 mm, UMZC Noy. 30, 1899;

c. Diogenes planimanus Henderson, 1893, 9 SL = 4.85 mm, UMZC

Nov. 30, 1899; d. Diogenes platvoeti nom. novy., holotypeo’ SL = 4.81

mm ZMA De201.872; e. Diogenes platvoeti nom. nov., 2 SL = 4.67

mm, UMZC Nov. 30, 1899; f. Diogenes mixtus Lanchester, 1902,

paralectotypeo’ SL = 5.81 mm, UMZC 1.10050.

DIOGENES OF SINGAPORE AND THE MALAY PENINSULA

margin denticulate over entire length. Dorsal margin of branchio-

stegite denticulate anteriorly and with 3 or 4 distinct spines posteriorly.

Ocular peduncles approximately 0.80 shield length, moderately

slender. Ocular acicles spinose along entire terminal margin. Inter-

calary rostriform process slightly overreaching tips of acicular

spines; with 2—5 spines on lateral margins in distal two-thirds and

terminal spine. Antennular and antennal peduncles approximately

equal in length, both overreaching ocular peduncles. Antennal acicle

weakly produced mesially, not distinctly forked, anterior margin

concave and spinulose or spinose.

Left cheliped (Fig. 12a) with upper margin of dactyl armed with

row of closely-spaced spinulose tubercles flanked on either side by

row of smaller tubercles; palm with 2 rows of moderately small

tuberculate spines on upper margin, proximal margin with row of

large blunt or spinulose tubercles; outer surfaces of dactyl, palm and

fixed finger all with blunt or acute tubercles, strongest in upper half

of palm, lower half of palm flattened, lower margin of palm and

fixed finger straight; carpus with double row of spines on upper

margin, outer face with irregular row of spines, strongest distally;

inner faces of palm and carpus tuberculate. Right cheliped with

spinulose upper margin of dactyl partially obscured by long setae;

upper margin of palm with irregular row of small spines, outer

surface granular or weakly tuberculate; carpus with row of strong

spines on dorsal margin and row of smaller spines centrally on outer

surface. Ambulatory legs with dorsal margins of dactyls each with

double row of small spines, mesial faces (Fig. 8c) each with longitu-

dinal row of small spines partially obscured by row of long setae;

propodi, carpi and meri each with double or triple rows of spines or

spinules on dorsal margins, lateral faces of propodi, carpi and meri

spinulose or tuberculate, distal margins of carpi also spinulose.

Males with paired gonopores, female with single right gonopore.

Telson (Fig. 9e) without distinct median cleft, but with markedly

asymmetrical lobes; terminal and lateral margins each with several

strong spines interspersed by small spinules.

COLOUR. Ocular peduncles, antennular and antennal peduncles

marked with alternating longitudinal stripes of cream and grey or

brownish grey. Rostrum and ocular acicles grey with tinge of red;

shield with dark grey-brown patches. Chelipeds and ambulatory legs

brown with dark brownish-grey patches (after Tirmizi and Siddiqui,

1982a).

DISTRIBUTION. Indian Ocean, including Bay of Bengal and north-

ern Arabian Sea; Malaysia; southeast coast of Australia.

REMARKS. Only one of the five syntypes is present in the BMNH’s

collection, i.e., a female, one of four specimens from Madras. The

fifth syntype is from Rameswaram. Presumably the remaining

syntypes are in the collection of the Indian Museum. Because of the

considerable morphological similarities among D. planimanus, D.

violaceus Henderson, 1893, D. intermedius De Man, 1892, and D.

custos (Fabricius, 1798), we are designating the female in the

BMNH collection (BMNH 1894:7:21:4) as the lectotype of D.

planimanus.

Lanchester reported eight specimens of D. planimanus, three

from ‘Loc. —?’ from Murex shells, and five from Patani from Natica

shells. The collection now consists of seven specimens: two (one

without a shell and one in a Thais sp. shell) presumably are those

from the unknown locality; five, of which four were still in shells of

two species of Natica, presumably are those from Patani. The five

latter specimens are indeed D. planimanus; the other two are not. Of

the specimens of D. planimanus, only one is complete, although its

left second and third pereopods are detached. The left chelipeds are

missing on the other four, as are most of the ambulatory legs. Despite

qd D C

d imm abcd

imm

1mm

egh

Fig.9 a. Telson, Diogenes goniochirus Forest, 1956, 9 SL = 2.15 mm,

UMZC Nov. 30, 1899; b. anterior lobe of sternite of 3rd pereopod,

Diogenes avarus Heller, 1865,0° SL = 1.53 mm, UMZC Nov. 30, 1899;

c. telson, Diogenes avarus Heller, 1865,0°SL = 1.53 mm, UMZC Nov.

30, 1899; d. telson, Diogenes avarus Heller, 1865,0 SL = 1.53 mm,

UMZC Nov. 30.1899; e, Diogenes planimanus Henderson, 1893, 9 SL =

4.85 mm, UMZC Nov. 30, 1899; f. telson, Diogenes platvoeti nom. nov.,

holotypeo’ SL = 4.81 mm ZMA De201.872; g. telson, Diogenes stenops

Morgan and Forest, 1991, ovigerous 9 SL = 3.11 mm, UMZC Nov. 30,

1899; h. telson, Diogenes platvoeti nom. nov., 2 SL = 4.67 mm, UMZC

Nov. 30, 1899; i. Diogenes mixtus Lanchester, 1902, lectotypeco’ SL =

6.88 mm, UMZC 1.10050.

some variation in the length and armature of the intercalary rostriform

process and relative lengths of the ocular, antennular and antennal

peduncles these specimens agree well with the lectotype.

Of the two remaining specimens, only one has appendages,

although the left cheliped is detached. These specimens are immedi-

ately distinguished from D. planimanus by the more triangular

shape of the shield, longer and more strongly armed intercalary

rostriform process, longer antennular peduncles, deeply forked

antennal acicles, and ambulatory dactyls that lack a row of spines on

the mesial faces.

Dechancé (1964) indicated that D. planimanus may have been

confounded with D. custos over a large portion of the range of the

latter. Diogenes custos, as described and illustrated by Tirmizi and

Siddiqui (1982a), does share some characters with Lanchester’s two

misidentified specimens, including the longer rostriform process

and unarmed mesial faces of the ambulatory dactyls. However, as

discussed below, Lanchester’s specimens represent D. intermedius.

The identity of specimens assigned toD. planimanus by Ajmal Khan

and Natarajan (1984: 16, fig. 13) is uncertain.

Diogenes platoveti nom. nov.

(Figs 5, 6, 8d, e, 9f, h, 12b)

Diogenes sp. De Man, 1892:352.

Diog. intermedius De Man, 1892:354.

Diogenes intermedius: Alcock, 1905:165 (list); Gordan, 1956:317

(list); Rahayu and Forest, 1995:385 (key), 387.

MATERIAL EXAMINED. Holotype by monotypy.c'(SL=4.81 mm),

Pare Pare, Celebes (Sudawesi, Indonesia), 1889, coll. M. Weber,

ZMA De.201.872; holotype of D. intermedius De Man, 1892.

Lanchester’s material. 1 9 (SL = 4.67 mm), 10°(SL = 5.86 mm)

lodged in shell of Thais sp., (Malaysia), collection site unknown,

‘Skeat’ Expedition; UMZC Nov. 30, 1899.

DIAGNOSIS. Shield (Figs 5, 6) slightly longer than broad, roundly

triangular; dorsal surface weakly spinulose and rugose; anterior

margin very weakly denticulate between obsolete rostrum and pro-

duced lateral projections. Dorsal margin of branchiostegite nearly

smooth, with sparse row of setae. Ocular peduncles approximately

Fig. 10 Left chela (outer face), a. Diogenes inglei sp. nov., holotype

ovigerous 9 SL = 1.46 mm, BMNH 1905.10.21.33; b. Diogenes

rectimanus Miers, 1884, holotypeco’ SL = 4.30 mm, BMNH 1882.7.Left

chela (outer face), a. Diogenes inglei sp. nov., holotype ovigerous 2 SL

= 1.46 mm, BMNH 1905.10.21.33; b. Diogenes rectimanus Miers,

1884, holotypeo’ SL = 4.30 mm, BMNH 1882.7.

P.A. MCLAUGHLIN AND P.F. CLARK

Fig. 11 Left chela (outer face), a. Diogenes goniochirus Forest, 1956, 9

SL = 2.15 mm, UMZC Nov. 30, 1899; b. Diogenes avarus Heller,

1865,0 SL = 1.53 mm, UMZC Nov. 30, 1899.

0.90 length of shield, moderately slender. Ocular acicles with 3 or 4

spines mesially and marginal row of very tiny spinules. Intercalary

rostriform process approximately 0.3 longer than longest acicular

spines; with 3—5 prominent spines, 4 or 5 additional much smaller

blunt spinules on lateral margins, and blunt or acute terminal spine.

Antennular peduncles overreaching antennal peduncles by 0.60—

0.75 length of ultimate segment, and ocular peduncles by entire

ultimate segment. Antennal acicle strongly bifurcate, outer projec-

tion slightly overreaching distal margin of penultimate segment,

inner reaching slightly beyond proximal half; anterior margins of

both spinose. Antennal flagellum with irregular long and short setae,

at least in proximal half.

Left cheliped (Fig. 12b) with upper margin of dactyl armed with

double row of closely-spaced small subacute spines, innermost

smallest, with intervening row of long setae; palm with double row

of larger subacute spines, proximal margin not distinctly delimited;

outer surfaces of dactyl, palm and fixed finger with scattered small

subacute or acute spines, largest forming faint arch medianly on

palm, lower margin of palm and fixed finger convex, with generally

double row of subacute spines; carpus with numerous small spines

on outer surface, strongest in lower half, upper margin with double

row of spines; inner faces of palm and carpus tuberculate. Right

cheliped with row of long stiff setae between and practically obscur-

ee ee

DIOGENES OF SINGAPORE AND THE MALAY PENINSULA

ing double row of small spines on upper margin of dactyl; upper

margin of palm with very short double row of small spines, outer

surface of palm, fixed finger and dactyl with widely scattered small

spines and tufts of long setae; carpus with row of spines on upper

margin, and outer face with scattered spinules and longitudinal row

of spines, all partially obscured by long setae. Ambulatory legs with

dorsal margins of dactyls each with somewhat irregular row of small

spines and long stiff setae, lateral surfaces each with longitudinal

sulcus, mesial surfaces (Fig. 8d, e) each with 2 longitudinal rows of

quite long setae; propodi each with double row of spines on dorsal

surface, carpi and meri each with single row of spines; lateral

surfaces of propodi and carpi tuberculate or spinulose, lateral faces

of meri nearly smooth; segments all with setose lateral surfaces.

Telson (Figs 9f, h) without median cleft, but with incipient lobes

markedly asymmetrical; smaller right lobe with few moderately

strong spines on terminal margin, lateral margin with several tuber-

cles or protuberances; elongate, subtriangular left lobe with few

spines on oblique terminal margin and series of spines on lateral

margin.

CoLour. Unknown.

DISTRIBUTION. Indonesia; Malaysia.

ETYMOLOGY. This species is named for Dirk Platvoet, Curator of

Crustacea, the Instituut voor Taxonomische ZoGdlogie, Zodlogisch

Museum, Universiteit van Amsterdam. The authors of this study

hope that honour has been fully satisfied and thank Dirk for his

persistence in locating the type of Diogenes intermedius De Man.

REMARKS. The species name Diogenes intermedius of De Man,

1892 is preoccupied by Diogenes pugilator var. intermedius Bouvier,

1891 (see page 404). According to ICZN 1985: 39, Article 16, a

name proposed with the term ‘variety’ or ‘form’ before 1961 does

not prevent availability [Art. 45g]. The species of De Man (1892) is,

therefore, given the replacement name Diogenes platvoeti nom. nov.

Of the two specimens in the Lanchester collection, only the

female has chelipeds and ambulatory legs. Both the holotype and the

Malaysian specimens are notable in lacking armature on the dorsal

margins of the branchiostegites. Lanchester’s specimens differ from

the holotype in having slightly longer antennular and antennal

peduncles. In these specimens the antennal peduncles overreach the

distal margin of the corneae by nearly the entire length of the

ultimate segment (Fig. 6); the antennal peduncles exceed the cor-

neae by 0.25-0.35 the length of the fifth segment. In the holotype,

the corneae are exceeded by only 0.75 the length of the ultimate

antennular segment (Fig. 6) and 0.15—0.20 the length of the fifth

segment of the antennal peduncles. De Man (1892) distinguished D.

intermedius {now D. platvoeti nom. nov.] from D. custos by the more

strongly bifurcate antennal acicles of his specimen. The acicles of

Lanchester’s specimens are similarly more strongly forked (Fig. 6),

differing from the holotype (Fig. 5) only in having the outer projec-

tion slightly broader and the inner projection slightly shorter. The

spination of the lateral margins of the second segment of the

antennal peduncle is slightly stronger in Lanchester’s material, but

the dorsal surface is more spinulose in the holotype. Minor differ-

ences between the holotype (Fig. 5) and Malaysian specimens (Fig.

6) have also been observed in the intercalary rostriform process,

which in the former is longer and more prominently spinulose in the

proximal half.

There is general agreement between Lanchester’s intact specimen

and De Man’s specimen as it pertains to armature of the chelipeds;

however, we did find a difference in the specific number of spines

present on the inner marginal row of the dactyl and on the upper

margin of the palm of the left cheliped. Similarly, the spines on the

outer face of the palm are somewhat stronger in the Malaysian

specimen than in the holotype. The row of spines on the dorsal

margin of each of the ambulatory dactyls is also stronger in

Lanchester’s specimens, and the median setal row on the mesial face

more complete (Figs 8d, e). The telsons of Lanchester’s specimens

are also more strongly armed, but the general configuration of the

lobes is comparable and the median cleft is absent in both (Figs 9f,

h). Tirmizi and Siddiqui (1982a) noted that females of D. custos have

a gonopore only on the right coxa of the third pereopods. Lanchester’s

female has paired gonopores, a character which distinguishes it not

only from D. custos, but also D. planimanus.

Diogenes stenops Morgan and Forest, 1991

(Figs 7a, 9g, 13b)

Diogenes senex: Lanchester, 1902:366; non Diogenes senex Heller,

1865.

Diogenes jousseaumei: Morgan, 1987b:179;non Diogenes jousseau-

mei (Bouvier, 1897).

Diogenes stenops Morgan and Forest, 1991:671, figs 9, 10.

MATERIAL EXAMINED. Paratypes: 20°(SL = 3.20, 5.40 mm), 1 2

(SL = 4.90), 25 mi south of Cairns, Queensland, 8 November 1965,

27 m, WAM 516-65; 1 oO (SL = 2.42 mm), New Year’s Island

Fig. 12

1893, 9 SL = 4.85 mm, UMZC Nov. 30, 1899; b. Diogenes platvoeti

nom. nov., 9 SL = 4.67 mm, UMZC Nov. 30, 1899.

Left chela (outer face), a. Diogenes planimanus Henderson,

Fig. 13. Left chela (outer face), a. Diogenes mixtus Lanchester, 1902,

paralectotypeo' SL = 5.81 mm, UMZC 1.10050; b. Diogenes stenops

Morgan and Forest, 1991, ovigerous 2 SL = 3.11 mm, UMZC Nov. 30,

1899,

(10° 54, 133° O1'E), October 1962, WAM 403-65. Lanchester col-

lection: 1 ovigerous ¢(SL = 3.11 mm), Pulau Bidan, Penang;

UMZC, Nov. 30, 1899.

DIAGNOSIS. Dorsal surface of shield (Fig. 7a) with tubercles and

spines, often in short transverse ridges. Ocular peduncles long and

slender, slightly overreached by antennular peduncles. Ocular acicles

with 3—5 spines on terminal margins. Intercalary rostriform process

very small, not reaching half length of ocular acicles. Antennal

peduncles slightly overreaching distal margins of corneae. Antennal

acicles with terminal spine and 3 or 4 spines on mesial margin.

Antennal flagella with long ventral setae.

Left cheliped (Fig. 13b) with dense plumose setae obscuring

armature, particularly on dactyl and palm; dactyl and palm with

row of strong spines on upper margin; outer faces of fixed finger

and palm with scattered tubercles or small spines, lower margins

with spines or spinulose tubercles; carpus with row of 6 or 7 very

strong spines on upper margin, distal margin with several spines, 1

or 2 very prominent spines on outer surface near distal midline.

Right cheliped with row of small spines on upper margin of dac-

tyl; upper margin of palm with strong distal spine and smaller

spines or tubercles proximally, outer face with slight to prominent

depression on outer face in upper half and scattered spinulose

P.A. MCLAUGHLIN AND P.F. CLARK

tubercles on outer surface of palm and fixed finger; carpus with

strong spine at upper distal angle; outer face with strong spine on

distal margin in upper half, outer surface and upper margin tuber-

culate or spinulose. Ambulatory legs with scattered long setae on

all segments; dactyls and propodi of second and right third un-

armed; carpi each with dorsodistal spine; left third slightly shorter

than right or second pereopods, ventral margin of propodus with

row of spinules; dactyl and propodus with appreciably more dense

tufts of setae, carpus with row of spinules or tubercles ventrolater-

ally and scattered tubercles on lateral surface, obscured by tufts of

setae.

Telson (Fig. 9g) with median cleft distinct, but not deep; posterior

lobes markedly asymmetrical, terminal margins with several large

and numerous smaller spines, extending onto lateral margins, at

least on left.

COLOUR. Shield cream and pale brown with darker patches. Ocu-

lar peduncles cream with some brown dorsally and ventrally; cornea

black with iridescent yellow speckling. Antennules and antennae

cream. Chelipeds cream and dark brown. Second and third pereopods

cream with grey-brown mottling, often with irregular brown band

proximally on dactyls and at mid-length of propodi, carpi and meri.

Setae pale grey, yellow or brown (after Morgan and Forest, 1991).

DISTRIBUTION. Northern Australia from the Northern Territory

east to the vicinity of Townsville, Queensland; Penang, Malaysia.

REMARKS. Morgan (1987b) reported Diogenes jousseaumei

(Bouvier), a species of the Troglopagurus group of Diogenes from

the Port Essington, Northern Territory, Australia, but after examin-

ing syntypic material of Bouvier’s (1897) species, concluded that his

Australian material represented a very similar, but specifically dis-

tinct taxon (Morgan and Forest, 1991). In this latter account, Morgan

and Forest commented that the previous record of D. jousseaumei

from Port Curtis, Queensland by Grant and McCulloch (1906)

should be regarded with “some suspicion’, and that the records of

this species from the Indian region (Alcock, 1905; Southwell, 1906)

might require substantiation. Haig and Ball (1988) reported D.

jousseaumei from the Arafura Sea and Torres Strait, and their colour

notes do not agree particularly well with those given by Morgan

(1987b) and Morgan and Forest (1991) for D. stenops, thus it is

possible that both species do occur in the region. Although actual

specimens were not in their collection, Rahayu and Forest (1995)

included D. jousseaumei in their key to Diogenes species in Indone-

sian waters; D. stenops was not mentioned from Indonesia, but 1n an

addendum, these authors reported its occurrence in Singapore.

Lanchester’s (1902) D. senex from Pulau Bidan, Penang, clearly

is not conspecific with Heller’s (1865) D. senex sensu stricto. The

markedly reduced intercalary rostriform process and heavy setation

of the cheliped unquestionably place it in the Troglopagurus group

of Diogenes. Despite the absence of the right cheliped, Lanchester’s

specimen compares very well with the four paratypes of D. stenops

we have examined. However, in the shape of the shield, Lanchester’s

specimen agrees better with their illustrated holotype (Morgan and

Forest, 1991, fig. 9) in having sloping anterolateral margins on the

shield. In all four paratypes, these margins are much straighter,

giving the shield a subquadrate appearance. The spination of the left

third pereopod of the Lanchester specimen corresponds quite well

with the smaller two paratypes. Apparently the spines on the ventral

margin of the carpus are reduced with increased animal size, as this

margin has only a row of minute, easily overlooked spinules in the

larger paratypes. The setation, particularly of the left chela, in

Lanchester’s specimen appears more coarse than any of the Austral-

ian paratypes. In the shape and armature of the antennal acicles,

DIOGENES OF SINGAPORE AND THE MALAY PENINSULA

Lanchester’s specimen does more closely resemble Morgan and

Forest’s (1991: fig. 11b) illustration of the syntype of D. jousseaumei;

however, it falls well within the range of variation in acicular length

and armature seen in D. stenops. It is quite possible that D. stenops

and D. jousseaumei may coexist geographically while occupying

different microhabitats. With the exception of shell occupancy

(Murex is reported for D. stenops), little is known about the ecology

of either species.

Another instance of geographic sympatry in species of the

Troglopagurus group has recently been documented. As previously

indicated, Diogenes stenops has been recorded from Singapore

(Rahayu and Forest, 1995), the type locality of D. jubatus Nobili,

1903. Although rarely reported and poorly known, the recent

redescription of D. jubatus by Lemaitre and Ng (1996) demonstrates

clearly the distinctiveness of the two species.

Diogenes mixtus Lanchester, 1902

(Figs 7f, 8f, 9i, 13a)

Diogenes mixtus Lanchester, 1902:367, pl. 34, figs 2, 2a, 2b; Nobili,

1903:16; Alcock, 1905b:165 (list).

MATERIAL EXAMINED. Lectotype (herein selected).c' (SL = 6.88

mm), Pulau Bidan, Penang, Malay Peninsula; UMZC I.10050.

Paralectotypes. 70°, 149, 1 ovigerous 9 (SL = 4.15-7.53 mm),

Pulau Bidan, Penang, Malay Peninsula, UMZC I.10050.

Additional material. 1 9 (SL = 4.95 mm), Kuching, Malaysia;

BMNH 1928.12.1.283.

DIAGNOSIS. Shield (Fig. 7b) with anterior margin weakly denticu-

late over 0.75 total length; rostrum obtuse, not reaching level of

lateral projections. Rostriform process elongate, overreaching ocu-

lar acicles by approximately third own length, multidenticulate.

Branchiostegites unarmed. Ocular peduncles overreached by both

antennular and antennal peduncles; ocular acicles broadly triangu-

lar, with 1 or 2 prominent spines mesially, with row of smaller spines

extending entire terminal margin. Antennal acicle bifurcate; mesial

fork with 0—2 small spines on outer margin and 2-4 smaller spines

on inner margin; lateral fork reaching beyond distal margin of fourth

peduncular segment, 4 or 5 small spines on inner margin and 0-3

spinules on outer margin.

Left cheliped (Fig. 13a) with outer face of palm armed with

double row of blunt spines extending from nearly proximal margin

almost to tip of fixed finger, with blunt spines scattered on fixed

finger and in somewhat irregular rows on ventral margin, midline of

palm with 2 irregular rows and tubercles dorsally; upper margin with

2 rather widely-spaced rows of small spines; dactyl with similar

rows of spines; carpus with double row of somewhat blunt spines on

upper surface, distal margin of outer face with row of spines curving

proximally near ventral margin; merus with acute row of spines on

dorsal margin, ventrolateral margin with row of acute spines,

ventromesial margin with double row of somewhat blunted spines.

Dorsal surface of palm of right cheliped with scattered spines,

partially obscured by long setae, dorsomesial margin with row of

small spines; upper outer and distal margins of carpus each with

irregular row of acute spines. Ambulatory legs generally similar;

dactyls (Fig. 8f) longer than propodi, dorsal margins with small

spines becoming obsolete in distal half; propodi each with 2 rows of

spines on dorsal surface; carpi each with row of spines on dorsal

margin.

Telson (Fig. 9i) with terminal margin minutely spinulose on right,

strongly spinose on left and approaching lateral angle, continued

onto lateral margin over approximately half length.

REDESCRIPTION. Shield (Fig. 7b) longer than broad, subovate to

subquadrate; anterior margin with row of closely-spaced small

tubercles or blunt spinules over 0.75 to entire width; rostrum obtuse

or broadly triangular, weakly produced, not reaching level of lateral

projections; lateral projections unarmed or with small to moderately

strong terminal blunt or acute spinule. Intercalary rostriform process

elongate, overreaching ocular acicles by 0.25—0.50 own length,

multidenticulate, with 3-7 lateral spines on each side of terminal

simple, bi- or trifid spine. Inner pterygostomial plate (cf. Pilgrim,

1973) with strong distal spine. Branchiostegites with upper margin

usually with row of closely-spaced small blunt or spinulose tuber-

cles partially obscured by tufts of long setae.

Ocular peduncles, moderately slender, corneae not particularly

dilated; overreached by both antennular and antennal peduncles.

Ocular acicles broadly triangular, usually with 1 or 2 more promi-

nent spines mesially, with row of smaller spines or spinulose tubercles

extending entire terminal margin.

Antennular peduncles overreaching ocular peduncles by 0.25 to

nearly entire length of ultimate segment; overreaching antennal

peduncles by 0.10—0.50 length of ultimate segment. Ultimate and

penultimate segments with scattered setae. Basal segment with row

of tiny spinules or tubercles on both distomesial and distolateral

margins.

Antennal peduncles with numerous long setae on fifth segment,

particularly dorsally and ventrally. Fourth segment with scattered

stiff setae. Third segment with spinule at ventrodistal angle. Second

segment with dorsolateral distal angle produced as acute spine,

lateral margin with low protuberances or spinules and long setae;

dorsomesial margin with row of small spinules, dorsal surface with

scattered spinules. First segment with row of tiny tubercles or

spinules on dorsal, dorsolateral and ventrolateral distal margins,

ventrodistal angle with strong spine. Antennal acicle strongly bifur-

cate; mesial fork with acute or bifid termination, 0—2 small spines on

outer margin and 2—5 smaller spines on inner margin; lateral fork

reaching to or beyond distal margin of fourth peduncular segment,

terminating in acute simple or bifid spine, 4-14 small spines on inner

margin and 0-4 spinules on outer margin, dorsal surface usually

with scattered spinules. Antennal flagellum moderately long, usu-

ally reaching to or beyond tip of left chela; several proximal articles

usually with 1 or 2 short or moderately long stiff setae; setae of

articles in distal two-thirds much shorter.

Left cheliped (Fig. 13a) with fingers opening nearly vertically;

cusp-like calcareous teeth on cutting edges of both dactyl and fixed

finger. Upper margin of dactyl armed with double row of small

spines and moderately dense, but relatively short setae; outer surface

with scattered small conical tubercles, row of tufts of stiff setae

adjacent to cutting edge; inner surface with scattered tufts of setae.

Palm with double row of subacute or acute spines on upper margin;

outer surface triangularly convex, with widely scattered conical,

often rather blunt spines on both lower half of palm and fixed finger,

2 irregular frequently rather widely-separated longitudinal rows of

slightly stronger spines in midline of palm and 1 shorter row in upper

half, row of blunt or subacute spines on lower margin, becoming

double row on fixed finger, lower and inner surfaces of palm

tuberculate. Carpus with double row of acute, subacute or blunt

spines on upper surface; outer face with numerous spines, strongest

in lower half, distal margin with row of small spines, lower margin

spinulose or spinose, inner face with weakly tuberculate or spinulose

distal margin, longitudinal row of spinulose protuberances or spines

and long setae near upper margin. Merus broadly triangular distally;

acute row of spines on dorsal margin, diverging distally into mesial,

dorsal and lateral rows extending to or nearly to distal margins;

dorso- and laterodistal margins with continuous row of moderately

strong, slender spines; ventrolateral margin with row of acute spines,

ventromesial margin with irregular double row of subacute or acute

spines, ventral surface spinulose or tuberculate. Ischium with spinules

or small spines on ventromesial and ventrolateral margins.

Right cheliped with moderately long and slender chela; fingers

opening nearly horizontally and terminating in strong calcareous

claws. Dorsal surface dactyl with 2 rows of spines and 1| additional

row on dorsomesial margin, all partially obscured by long setae. Palm

with scattered moderately strong spines, partially obscured by long

setae, dorsomesial margin with single or row of small spines, mesial

face somewhat spinulose; fixed finger with 2 or 3 rows of small

spinules and tufts of long setae on dorsal surface; dorsolateral margin

not well defined, but with numerous small spinules. Carpus broadly

triangular; dorsomesial margin with row of small spines practically

obscured by long dense setae, and adjacent row of stronger spines on

dorsal surface, dorsolateral margin with single or irregular double

row of spines and tufts of setae, distal margin with several spines;

lateral face spinulose; mesial face weakly tuberculate. Merus triangu-

lar; dorsal margin with row of spinules or small spines and tufts of

long setae, 2 or 3 prominent spines at or near distal margin, often |

additional strong spine marginally just laterad of midline; dorso- and

laterodistal margins usually with small spinules; lateral face fre-

quently with numerous short multifid ridges; ventrolateral margin

with row of strong spines or acute spines distally and small, multifid

short ridges proximally; ventromesial margin with generally double

row of small spinules. Ischium with row of small spines or spinules on

ventromesial margin; laterodistal margin with few spinules.

Ambulatory legs generally similar form left to right; dactyls (Fig.

8f) long, approximately 0.20 longer than propodi, slender, curved,

slightly twisted; ventral margins each with row of long setae; lateral

faces each with longitudinal suture; dorsal margins with small

spines becoming obsolete in distal half and long setae; mesial faces

each with longitudinal sulcus lined with long setae, row of small

spines ventrally decreasing in size and not reaching to distal third

and gradually replaced by row of long setae. Propodi each with 2

rows of spines on dorsal surface, strongest mesially and separated by

flattened, unarmed or intermittently spined longitudinal space; lat-

eral face with longitudinal row of spinulose tubercles or spines

dorsally and usually numerous simple or multidenticulate tubercles,

sometimes only weakly developed; ventral surface faintly spinulose,

ventrodistal margin usually with row of denticles extending mesially

and laterally; mesial face somewhat spinulose ventrally or with

irregular longitudinal rows of small tubercles. Carpi each with row

of strong spines on dorsal margin; lateral face with 3-5 usually

longitudinal rows of spines, spinules, multidenticulate tubercles or

low protuberances; laterodistal margin spinose or spinulose; mesial

faces each with longitudinal row of small spines adjacent to dorsal

margin (second) or unarmed (third). Meri with dorsal surfaces of

second pair distally broadened and armed with irregular double or

triple rows of small spines or spinulose tubercles, dorsal margin

proximally and on third pereopods each with row of spines;

ventromesial margins each with nearly double row of spines or

spinulose tubercles; row of small spines on ventrolateral margins;

lateral faces, particularly of third weakly spinulose or tuberculate.

Sternite of third pereopods subrectangular, with tuft of setae on

either side laterally. Sternite of fifth pereopods as slender elongate,

tuberculate, calcareous rod.

Protopod of right uropod with well developed posterior protuber-

ance nearly equal to size of endopod, and similarly covered with rasp

of corneous scales. Telson (Fig. 91) without median cleft; terminal

margin spinulose on right, strongly spinose approaching left lateral

angle and continuing onto lateral margin over approximately half

length.

P.A. MCLAUGHLIN AND P.F. CLARK

COLOUR. Not known.

DISTRIBUTION. Malaysia; ? Singapore.

REMARKS. Nobili (1903) reported numerous specimens of D.

mixtus collected in Singapore. He distinguished Lanchester’s (1902)

species from D. intermedius De Man, 1892 by the presence, in the

former species, of spines on the merus of the second and third

pereopods. A check of the collections of the Museo Regionale di

Scienze Naturali, Torino, failed to locate Nobili’s specimens (Elena

Gavetti, pers. comm.); therefore it has not been possible to confirm

Nobili’s (1903) identification. Although it was not represented in

their collection, D. intermedius was reported as one of the Indone-

sian species of Diogenes by Rahayu and Forest (1995); no mention

was made of D. mixtus.

Paguristes hians Henderson, 1888

(Fig. 14)

Paguristes hians Henderson, 1888:79, p. 8, fig. 4; Alcock, 1905: 40,

pl. 3, fig. 2; Southwell, 1906:216; Estampador, 1937:506;

Thompson, 1943:415; Gordan, 1956:322 (in part) (lit.); Edwards

and Emberton, 1980:236 (list); Haig and Ball, 1988:173; Hogarth,

1988:1100; Morgan, 1990:21.

Diogenes desipiens Lanchester, 1902:366, pl. 34, figs 1, 1a; Alcock,

1905:165 (list); Gordan, 1956:317 (lit.).

Non Paguristes hians: Grant and McCulloch, 1906:33; McCulloch,

1913:346 = Paguristes monoporus Morgan, 1987.

MATERIAL EXAMINED. Holotype. O(SL = 4.89 mm); BMNH

1888.33., H.M.S. ‘Challenger’ station 208, off Manila. Supplemen-

tal material. | Oo’, 2 ovigerous 9 (SL = 1.67—2.51 mm), Maldive

Islands, 14 February, 20 December 1993, collector P. Hogarth.

DIAGNOSIS. Shield (Fig. 14) considerably longer than broad. Ros-

trum broadly triangular, not produced to level of terminal spinules of

obtusely triangular lateral projections. Ocular peduncles long and

extremely tenuous, slightly overreaching antennular peduncles and

approximately 0.50 longer than antennal peduncles; acicles elongate

and nearly rectangular, with large tuberculate terminal spine and

smaller accessory spine laterally. Subquadrate calcified lobe with 4

subacute spinules on anterior marginal part of, or articulating with,

interocular lobes. Antennal acicle long, reaching nearly to mid-

length of ultimate peduncular segment.

Fig. 14 Paguristes hians Henderson, 1888, holotypeo' SL = 4.89 mm,

BMNH 1888.33, a. whole animal; b. enlarged view of shield and

cephalic appendages.

DIOGENES OF SINGAPORE AND THE MALAY PENINSULA

Chelipeds (Fig. 14) similar, subequal; dactyls and fixed fingers

with distinct hiatus; dorsal surfaces of palms relatively smooth

proximally with only few spinules, stronger spines distally; dorsal

surfaces of carpi each with irregular row of spinules becoming

strong spines at distal margin. Ambulatory legs with dactyls slightly

longer than propodi; propodi and carpi of second each with row of

spines on dorsal margins, partially obscured by long setae; propodi

of third unarmed, carpi each with spine on dorsodistal margin.

Males lacking paired second pleopods; first paired, but consisting

of short, broad, uniramous 2-segmented appendages. Telson with

posterior lobes nearly symmetrical; terminal margins rounded, armed

with 5 or 6 spines.

REDESCRIPTION. Shield (Fig. 14) subtriangular, considerably

longer than broad, with numerous small spinules, spinulose tuber-

cles and tufts of plumose setae on dorsal surface, particularly

laterally. Rostrum broadly triangular, with very small terminal

spinule, not produced to level of terminal spinules of obtusely

triangular lateral projections. Branchiostegite with 4 or 5 slender

acute spines on dorsodistal margin partially to entirely obscured by

long setae.

Ocular peduncles long and slender, slightly overreaching

antennular peduncles and approximately 0.50 longer than antennal

peduncles, dorsomesial surface with row of long setae; corneae

small, not dilated. Ocular acicles elongate and nearly rectangular,

dorsally flattened, with large tuberculate terminal spine and smaller

accessory spine laterally, with several long plumose setae distally.

Subquadrate calcified lobe (with 4 subacute spinules anteriorly in

holotype) seemingly articulated with interocular lobes.

Antennular peduncles not reaching to bases of corneae; ultimate

and penultimate segments unarmed; basal segment with very strong

curved spine on dorsolateral distal margin, smaller spine on

ventrodistal margin.

Antennal peduncles with supernumerary segmentation; reaching

only to about distal third of ocular peduncles; fifth, fourth and third

segments each with strong ventrodistal spine and scattered long

setae, most abundant laterally; second segment with dorsolateral

distal angle produced, terminating in bifid spine, lateral margin with

row of long setae, dorsomesial distal angle with strong spine; first

segment unarmed but with long setae laterally. Antennal acicle long,

reaching nearly to mid-length of ultimate peduncular segment,

broad and dorsally flattened, lateral margins each with 2 or 3 strong

spines and row of very long plumose setae, mesial margins each also

with 2 or 3 strong spines and row of plumose setae sufficiently long

to form a setal net above antennules, terminating in bifid spine.

Antennal flagellum short, not reaching beyond proximal margins of

chelipeds; each article with 1 or 2 long and 1 or 2 short setae.

Chelipeds (Fig. 14) similar, subequal, right slightly larger (at least

in males). Dactyls approximately twice length of palms; dorsomesial

margins each with row of spines, dorsal and mesial surfaces with

fairly closely-spaced tuberculate spines and tufts of long setae;

cutting edges each with 1 or 2 large calcareous teeth proximally and

row of smaller calcareous teeth in distal 0.66, terminating in cor-

neous claw; dactyls and fixed fingers ventrally curved and with

distinct hiatus. Palms shorter than carpi, dorsomesial margins each

with row of 2 to 4 strong conical spines, dorsal surface relatively

smooth proximally with few spinules or spinulose tubercles, stronger

spines distally tending to form 1 or 2 irregular rows, and on weakly

delimited lateroproximal margin; fixed finger with dorsolateral

margin not clearly delimited, dorsal and lateral faces with closely-

spaced spinulose tubercles and spines; cutting edge with row of

calcareous teeth, strongest proximally; ventral surface of palm with

few scattered tubercles; all surfaces with tufts of long setae. Carpi

trapezoidal in dorsal view; slightly less than 0.50 length of meri;

dorsomesial margins each with row of 4 strong conical spines, first

1-3 with very small spinule basally on mesial side, dorsolateral

margins each with irregular row of smaller spines, dorsal surface

with irregular row of spinules becoming strong spines at distal

margin; ventrolateral distal margins each with | or 2 small tubercles;

surfaces with long setae. Meri each with longitudinal row of spinules

on dorsal margin in proximal 0.65, distally 1 transverse row of

prominent spines extending onto lateral and mesial faces dorsally

and second similar row on distal margin; ventromesial margin with

row of acute spines; ventrolateral margin with row of small spines in

distal half and short transverse rows of spinules proximally. Ischia

each with row of spinules on ventromesial margin and 1 small spine

at ventrolateral distal angle.

Ambulatory legs with dactyls slightly longer than propodi; in

dorsal view slightly twisted, in lateral view curved ventrally in distal

halves; all surfaces, and particularly dorsal and ventral margins, with

rows of long setae. Propodi of second each with row of spines on

dorsal margins; carpi with single or double row of spines, all

partially to completely obscured by long setae; propodi of third (left

third broken at distal margin of ischium in holotype) unarmed but

with tufts of long setae on all surfaces, carpi only with spine at

dorsodistal margin or with 2 or 3 small spines in distal half, all at

least partially obscured by tufts of long setae. Meri of second each

with row of very small spinules and tufts of long setae on dorsal

margins, ventral margins each with row of spines and tufts of long

setae; third with few minute spinules on dorsal margin and tufts of

long setae, ventral margin with tufts of long setae. Ischia each with

1 or 2 spinules on dorsal margins, ventral margins each with row of

long setae and | small spine near distal margin.

Males with paired gonopores; no paired second pleopods; first

paired but consisting of short, broad, uniramous 2- segmented

appendages positioned directly over coxae of fifth pereopods; un-

paired left pleopods 3 to 5 uniramous. Females with single left

gonopore; paired first pleopods modified as gonopods; very large

brood pouch. Telson with prominent, deep transverse suture; poste-

rior lobes nearly symmetrical, terminal margins rounded, each

armed with 4 to 6 spines, some corneous-tipped, and few moderately

long setae.

COLouR. Not reported.

DISTRIBUTION.

sia; Philippines.

Oman; Maldives; Red and Arabian Seas; Malay-

REMARKS. Henderson (1888) described the species from a single

male; however, he made no comment on any of the abdominal

appendages. Although Paguristes hians is a very distinctive species

that possesses characters unique among species of Paguristes, only

the remarks by Haig and Ball (1988) called attention to any of these.

These authors appear to have been the first to document that males

lacked the paired second pleopods typical of species of Paguristes;

they also pointed out the presence of a single left gonopore in the

female. It may be that these abnormalities were recognized by Grant

and McCulloch (1906) and McCulloch (1913) who incorrectly

identified atypical Paguristes specimens from Mast Head Island,

Queensland, Australia, as P. hians. Haig (in Haig and Ball, 1988)

showed that at least some of the Mast Head Island specimens

actually were referable to P. monoporus Morgan. As described by

Morgan (1987a) this is another rather bizarre species of Paguristes

in which male first and second pleopods are reduced and restricted

to the right side of the abdomen; males possess only a single right

gonopore and females have only a single left.

While the characters described above for P. hians set this species

apart from all other known Paguristes species, they are not the

characters that drew our attention to the similarities between this

species and Lanchester’s (1902) Diogenes desipiens. Paguristes

hians has uniramous unpaired male pleopods, a character com-

monly associated with species of Diogenes. Similarly, P. hians has

a subquadrate calcified lobe that is, at least positionally, similar to

the intercalary rostriform process that distinguishes most Diogenes

species from other Diogenidae. In the holotype of P. hians this

structure is armed with 4 marginal spinules. In the three small

specimens from the Maldive Islands, this process appears almost

identical to that of D. desipiens as shown by Lanchester (1902: fig.

1); in one specimen, it appears denticulate under high magnifica-

tion, as described for D. desipiens. Additionally, the distinctive

shape, armature, and setation of the ocular and antennal acicles of

P. hians are virtually identical to those described and illustrated by

Lanchester for D. desipiens. Lanchester’s description of the shield

armature, as well as ocular peduncle length and its relationship to

the lengths of both the antennular and antennal peduncles agrees

extremely well with those of P hians. Furthermore, Lanchester

described the chelipeds of his species as being subequal, the right

being slightly larger. We know of no Diogenes species in which the

chelipeds are subequal, but P hians has subequal chelipeds, the

right of which is slightly larger in the holotype and male specimen

from the Maldive Islands. Lanchester’s description of the armature

of the chelipeds also agrees quite closely with the type of P. hians

and the three smaller specimens that we examined. Similarly there

is agreement between the length ratios of the dactyls and propodi

of the ambulatory legs of the two species. However, disagreement

between Lanchester’s description of D. desipiens and our observa-

tions of P. hians is found in the armature of these appendages. We

observed that the ventral margins of the dactyls of the holotype

have a row of slight protuberances; the Maldive Islands specimens

each have a row of corneous spinules on these margins. The dorsal

margins of the propodi and carpi of the second pereopods each

carries a row of spines, and the carpi of the third may have from |

to 3 spines on the dorsal margins in our specimens. The dorsal

margins of the meri of both pairs of pereopods are armed with

spines; the ventral margins of the second pereopods also are

spinose. But all of the spines are at least partially obscured by tufts

of setae. Lanchester described the ambulatory legs as being

‘densely hairy on their upper and lower margins, otherwise

smooth.’ Whether Lanchester simply failed to notice spines

amongst the setal tufts on his specimen, or they were actually

lacking, is a matter of speculation. Given all of the other similari-

ties between the two taxa, we are inclined to presume the former.

Therefore, until specimens having all of the attributes accredited to

Lanchester’s taxon are found in a species unquestionably referable

to Diogenes, we consider D. desipiens a junior subjective synonym

of Paguristes hians.

ACKNOWLEDGEMENTS. We express our gratitude to Dr. Raymond

Symonds, University Museum of Zoology, Cambridge, for the loan of

Lanchester’s ‘Skeat’ Expedition material and providing supplemental infor-

mation on the collection including Diogenes desipiens; Dr. Peter Ng, National

University of Singapore for search the Institute reference collections in an

unsuccessful attempt to locate the type of Diogenes desipiens; Dr. Peter

Hogarth, Biology Department, University of York for permitting us to use his

specimens of Paguristes hians from the Maldive Islands; Dr. Diana Jones,

Western Australian Museum for the loan of paratypes of Diogenes stenops;

Drs L.B. Holthuis and Michel Tiirkay for information concerning the wherea-

bouts of the Holy Grail, i.e., Diogenes intermedius; and Drs F. Schram and

Dirk Platvoet, Instituut voor Taxonomische Zodlogie (Zodlogisch Museum),

Universiteit van Amsterdam, for the loan of De Man’s Diogenes intermedius.

P.A. MCLAUGHLIN AND P.F. CLARK

The photographs are the work of the late E.J. McGeorge. The assistance of

George Holm, Richmond, British Columbia, Canada, and Janet Armbrust,

Mt. Vernon, Washington, in the identification of Lanchester’s shells is ac-

knowledged. This is a scientific contribution from the Shannon Point Marine

Center, Western Washington University.

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Issued 26 June 1997

A new species of crassispirine gastropod from the

Houtman Abrolhos Islands, Western Australia

(Gastropoda, Conoidea, Crassispirinae)

ALEXANDER V. SYSOEV

Zoological Museum of Moscow State University, Herzen Street 6, Moscow 103009, Russia

JOHN D. TAYLOR

Department of Zoology, The Natural History Museum, London SW7 5BD, U.K.

SYNOPSIS.

A new species of conoidean gastropod, Burchia spectabilis, is described from the Houtman Abrolhos Islands,

Western Australia. Although the wishbone-type radular teeth are similar to those of Inquisitor, the shell is unlike any Indo-Pacific

species and most similar to the West American genus Burchia.

INTRODUCTION

In this paper we describe an distinctive new species of crassispirine

gastropod dredged from around the Houtman Abrolhos Islands,

Western Australia. Unfortunately, we have only two specimens of

this species, one of which was serial sectioned to determine the

anatomy of the foregut which is described in the following paper

(Kantor, Medinskaya & Taylor 1997).

SYSTEMATIC DESCRIPTION

Family TURRIDAE H. & A. Adams, 1853 (1840)

Subfamily Crassispirinae McLean, 1971

Burchia spectabilis Sysoev & Taylor, new species

Figs 1-3

TYPE MATERIAL. Holotype: Western Australian Museum, Perth,

WAM 123-9. Paratype: serial sections of the foregut, Natural His-

tory Museum, London.

TYPELOCALITY. Goss Passage, Wallabi Group, Houtman Abrolhos

Islands, Western Australia (Glover & Taylor, 1997: station 11),

28°28.64' S, 113°46.45' E, depth 41m, coarse sand and gravel.

DIAGNOsIS. Shell rather large and thick-walled, fusiform, with

high and many-whorled spire occupying about half of the shell

height. Shell colour pinkish white, with darker reddish-brown

siphonal canal and some interspaces between sculptural elements,

and blotches of the same colour on the subsutural ramp. Protoconch

consists of 1.7 smooth whorls. Initial teleoconch whorls almost flat-

sided, then angulate at the periphery. Subsutural fold strong and

tuberculate. Subsutural ramp concave and devoid of axial sculpture.

Axial sculpture of numerous folds forming low and blunt tubercles

below the subsutural ramp. Folds usually branch towards the lower

suture. On last teleoconch whorls the folds become irregular. Spiral

sculpture of deeply incised and widely spaced grooves becoming

closer to each other on the canal. Aperture rather narrow. Siphonal

canal short and wide, with distinct notch. Parietal nodule moderately

developed. Anal sinus deep, U-shaped, with the apex in the middle

of subsutural ramp. Operculum large and thick, oval, with a terminal

nucleus. Radula of wishbone marginal teeth only, teeth rather nar-

row, with moderately wide accessory limb.

DESCRIPTION OF HOLOTYPE. The shell is rather large, strong,

fusiform, with high spire comprising 0.47 of the shell height. The

ground shell colour is pinkish-white, with pale brownish spiral band

just below the body whorl periphery, reddish-brown siphonal canal

and the lower part of the shell base, and some interspaces between

sculptural elements, and occasional blotches of the same colour on

the subsutural ramp. The periostracum is poorly developed over the

entire shell except for the lower part of the shell base and the canal.

The protoconch is rather pupiform and consists of 1.7 smooth glossy

whorls with shallow suture. The border between protoconch and

teleoconch is not clearly defined. The teleoconch consists of 10

whorls separated by rather shallow and slightly wavy sutures. Initial

teleoconch whorls are almost flat, then a peripheral angulation

appears, which becomes stronger towards the body whorl The

subsutural fold is well developed and tuberculate. The subsutural

ramp is rather deeply concave. The whole shell surface is wrinkled

by numerous rough growth lines. The axial sculpture consists of

numerous folds beginning below the subsutural ramp and forming

wide, low and blunt tubercles. Below the tubercles, the folds usually

branch into two or three narrower folds separated by narrow

interspaces. Sometimes additional folds appear below the row of

tubercles and widen towards the lower suture. On the body whorl the

folds become even more irregular and of varying width; they may

branch or merge along their length. The folds reach the siphonal

canal. There are 13 tubercles below the subsutural ramp on the

penultimate whorl, and about 18 on the body. The spiral sculpture is

represented by incised and widely spaced narrow grooves. There is

one such groove below the suture, 4-5 more crowded on the subsutural

ramp, and 3 below it on teleoconch whorls. On the body whorl, there

are 3 grooves below the subsutural ramp and, after a very wide

interspace at the periphery, 3 grooves on the shell base followed

by 15 closely spaced ones on the canal. The last quarter of the body

whorl is marked by a strong and wide varix, which has no clearly

defined borders and looks more like a longitudinal swelling.

The aperture is rather narrow, more or less uniform in width,

including the short and wide canal with a distinct notch The

columellar part of aperture is almost straight and covered with a

A.V. SYSOEV AND J.D. TAYLOR

Fig. 1 Burchia spectabilis Sysoev & Taylor. Holotype. Shell height 36.9 mm.

Fig. 2 Holotype, lateral view

Fig. 3 Burchia spectabilis, holotype, radula teeth. Scale bar = 20ym.

thick, white callus. The parietal nodule is moderately developed.

The anal sinus is deep, U-shaped, broadly open, symmetrical, with

the apex in the middle of subsutural ramp. The operculum is large

and thick, oval, dark-brown, with a terminal nucleus, 7.2 x 3.2 mm.

Shell height is 36.9 mm, body whorl height 19.5 mm, aperture height

14.4 mm, and shell diameter 10.4 mm. Paratype: shell height 12.5

mm.

RADULA. The radula (Fig. 3) consists of two rows of wishbone

type marginal teeth only. The teeth are rather narrow, slightly curved

and sharply pointed, without barb or cutting edge. The accessory

limb is moderately large and attaches to the major limb about just

behind the tip.

ANATOMY. The foregut anatomy is described in Kantor et al. 1997

(p. 81)

REMARKS. On shell characters, it is difficult to assign this species

to any genus of Crassispirinae known from Western Australia and

the entire Indo-Pacific. Its radular teeth are very similar to those of

species of Inquisitor Hedley, 1918 (see Taylor & Wells, 1995). In

general shell outline the new species resembles some species in-

cluded by Wells (1994) in the genus Inquisitor Hedley, 1918 namely

I. dampierius (Hedley, 1922) and, to a lesser extent, /. odhneri Wells,

1994, but is readily distinguished from them (and from any known

turrid species) by the peculiar character of the sculpture of branching

axial folds and widely spaced spiral grooves. However, the two latter

species are themselves not very similar to the type-species of

Inquisitor, I. sterrhus (Watson, 1881), and other typical representa-

tives of the genus, which are characterized by a slender shell with a

well differentiated and relatively long siphonal canal. Unfortunately,

the radular characters of J. dampierius and I. odhneri are unknown.

On the other hand, the radular and conchological characters of the

new species correspond quite well to those of Burchia Bartsch,

1944, a Central American subgenus of Crassispira Swainson, 1840.

The main feature distinguishing species of Burchia from the new

species, besides the unique sculpture of the latter, is the presence of

a thick periostracum. However, the character of periostracum is not

usually considered to be of taxonomic importance at the generic

level. Therefore, it seems reasonable under these circumstances to

place the new species, at least provisionally, into Burchia.

ACKNOWLEDGEMENTS. This species was collected during the Interna-

tional Marine Biological Workshop on the Marine Fauna and Flora of the

Houtman Abolhos Islands. We are grateful to the organiser Dr Fred Wells of

the Western Australian Museum and the crew of the fisheries research vessesl

‘Flinders’ for facilities and logistic support.

A NEW SPECIES OF CRASSISPIRINE GASTROPOD

REFERENCES

Glover, E.A. & Taylor, J.D. 1997 in press. Diversity and distribution of subtidal

molluscs from the outer continental shelf, Houtman Abrolhos Islands, Western

Australia. Jn: Wells, F.E. (ed.) The Marine Fauna and Flora of the HoutmanAbrolhos

Islands, Western Australia. The Western Australian Museum, Perth.

Kantor, Y.I., Medinskaya, A. & Taylor, J.D. 1997. Foregut anatomy and relationships

of the Crassispirinae (Gastropoda: Conoidea). Bulletin of the Natural History

Museum, London (Zoology) 63: 55-92.

38)

Taylor, J.D. & Wells, FE. 1994. A revision of the crassispirine gastropods from Hong

Kong (Gastropoda: Turridae). pp. 101-116. Jn: Morton, B. (ed.)The Malacofauna of

Hong Kong and Southern China III. Proceedings of the Third International Work-

shop on the Malacofauna of Hong Kong and Southern China, Hong Kong University

Press, Hong Kong.

Wells, FE. 1994. A revision of the Recent Australian species of the turrid genera

Inquisitor and Ptychobela. Molluscan Research 15: 71-102.

Bull. nat. Hist. Mus. Lond. (Zool.) 63(1): 55—92

Foregut anatomy and relationships of the

Crassispirinae (Gastropoda, Conoidea)

YURI I. KANTOR AND ALEXANDRA MEDINSKAYA

A.N. Severtzov Institute of Animal Evolutionary Morphology and Ecology, Russian Academy of Sciences, Lenin

Avenue 33, Moscow 117071, Russia

JOHN D. TAYLOR

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK

CONTENTS

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SYNOPSIS. The foregut anatomy of 31 species from the conoidean subfamily Crassispirinae is described. Great variation is

found between species in the configuration of the foregut including features such as: the structure of the rhynchodeum, the

morphology of the proboscis, the position and number of buccal tube sphincters, the position and structure of the buccal mass,

the histology of the salivary glands and venom gland, the length of the oesophagus, the structure of the muscular bulb and the

morphology of the radular teeth. Many species have marginal teeth of the wishbone type, but teeth are paddle-shaped in Funa and

Vexitomina, harpoon-shaped in Cheungbeia and a hollow awl shape in Ptychobela. Many crassispirinae have rather similar shells,

but different anatomies. Some species with similar radulae have different anatomies and others with similar anatomy have widely

differing radulae. An analysis of relationships, using Gemmula as outgroup, shows that the various subgenera of Crassispira are

not monophyletic and should be raised to generic status. Also, Epidirona is polyphyletic with some species belonging in the

Turrinae, as do some species of Turridrupa. Additionally, some species classified in the genera Guraleus, Antiguraleus and

Issued 26 June 1997

Paraguraleus, until recently classified as Mangeliinae, belong in the Crassispirinae.

INTRODUCTION

Gastropods of the superfamily Conoidea are notable for the posses-

sion of a large, coiled venom gland, together with the highly

modified radular teeth which are used to inject the venom into the

prey. Although Conus is the most well-known taxon, it represents

only a small part of the total diversity of the group which conserva-

tive estimates suggest as more than 4000 living species and 340

genera (Taylor, Kantor & Sysoev, 1993). Although most classifica-

tions (e.g. Powell, 1966; McLean, 1971) have been based largely on

shell and radular characters, Taylor et al. (1993) have recently

provided anatomical criteria, mainly derived from characters of the

foregut, for the definition of suprageneric taxa of conoideans. Al-

though their study involved anatomical investigation by serial sections

of more than 72 species of conoideans, this nevertheless represented

only a small fraction of the living genera and species. Moreover,

amongst the species studied so far, a wide disparity in the configura-

tion of the various organs of the foregut in the Conoidea has been

revealed (Taylor et al., 1993), with new arrangements still being

discovered (Kantor & Taylor, 1994;Taylor, 1994; Kantor and Sysoev,

1996). These preliminary studies suggested that the subfamily

Crassispirinae, one of four subfamilies of Turridae possessing “wish-

bone’ radular teeth, showed a wide variation in radular morphology

and foregut anatomy, including the possibility of further evolution-

ary pathways to the hypodermic feeding system. Moreover, within

the Crassispirinae, some species in the genera /nguisitor, Funa and

Ptychobela which possess rather similar shells, were shown to have

very different foregut and radular morphologies (Kilburn, 1989;

Taylor, 1994; Taylor & Wells, 1994). The problem of using shell

characters alone to classify conoideans has recently been high-

lighted in the case of the southern African species Antiguraleus

morgani, previously classified in the Mangeliinae on the basis of

shell morphology, but shown to be a likely crassispirinan on radular

characters (Kilburn, 1994). For these reasons we decided to investi-

gate the anatomy of the Crassispirinae in more detail.

Currently, some 48 genera and subgenera have been assigned to

the Crassispirinae (Taylor et al., 1993; Taylor & Wells, 1994;

Kilburn, 1994) mainly on the evidence of radular characters, but in

some cases on shell morphology alone. The subfamily is diverse in

the tropical West America (McLean, 1971; Keen, 1971) with over 52

species recorded, and also in the Caribbean (Maes, 1983) and West

Africa (Fernandes, Rolan & Otero-Schmitt, 1995). The Indo-Pacific

fauna is less well known and there are many undescribed species, but

Kilburn (1988; 1994) reports 54 species from Southern Africa,

including numerous new genera and species, and many other species

Table 1 List of crassispirine gastropods and the outgroup Gemmula

which have been sectioned, with details of their collection locations.

Crassispira (Crassispira) incrassata (Sowerby, 1834). North side of

Venado Island, Panama, (8°53’N, 79°36’ W). ANSP A9695 T357.

Crassispira (Crassispira) maura (Sowerby, 1834). Between Tortota and

Venado Island, Bay of Panama, Republic of Panama (8°51°45N,

79°35°40W), 9-10 m, ANSP A9613 T357.

Crassispira (Gibbaspira) dysoni (Reeve, 1846). Isla de Lobos, Gulf of

Mexico, Mexico (21°28’N, 97°13’ W). ANSP A9423 T357.

Crassispira (Glossispira) harfordiana flucki (Brown & Pilsbry, 1913).

Playa Benito, Campeche, Gulf of Mexico, Mexico (19°48’N, 90°36’ W).

ANSP 356707 A9734 T1357.

Crassispira (?Crassispirella) latizonata (E.A. Smith, 1882). 1 km north of

Holotown, St James, Barbados (13°11’N, 59°38’ W). ANSP A9866G

WS.

Crassispira (Monilispira) pluto Pilsbry & Lowe, 1932. West San Carlos

Bay, Sonora, Mexico (27°57’N, 111°04’°W). ANSP A9226 T357.

Crassispira (Striospira) kluthi Jordan, 1936. Venado Island, Panama

(08°53’N, 79°36’ W). ANSP 356323 A9698 T357.

Crassispira (Striospira) tepocana Dall, 1919. San Carlos, Guaymas,

Sonora, Mexico, 35 m. ANSP A6670.

Crassispira (Striospira) xanti Hertlein & Strong, 1951. West side of

Viradores sur, Bahia del Cocos, Costa Rica (10°34’45N, 85°43’35W).

ANSP 35791A10167 T357.

Crassispira (Crassiclava) turricula (Sowerby, 1834). Off Nacascola, west

side of Bahia Culebra, Bahia Culebra, Costa Rica (10°37°15N,

85°41’20W). ANSP A9753B T357.

Crassispira (Crassiclava) apicata (Reeve, 1845). Cactus Point, Prickly

Pear Island, British Virgin Islands (18°30°55N, 64°22’30W). ANSP

355532 A9461E T357.

Burchia spectabilis Sysoev & Taylor, 1997. Houtman Abrolhos Islands,

Western Australia (see Sysoev & Taylor, 1997). BM(NH).

Miraclathurella bicanalifera (Sowerby, 1834). West side of Viradores Sur,

Bahia del Cocos, Costa Rica (10°34.30’N, 85°43.40’W). ANSP 357806

A9857A T357.

Hindsiclava andromeda (Dall, 1919). 3 miles southwest of Punta San

Antonio, Sonora, Gulf of California, Mexico (27°54’N, 111°08’W).

ANSP 358149 A10192G T357.

Hindsiclava militaris (Reeve, 1843) 2 miles east of Punta Doble, Sonora,

Gulf of California, Mexico (27°55’N, 111°04’W). ANSP A10186 T357.

West side of Bahia Culebra, off Nacascola, Bahia Culebra, Costa Rica

(10°37’ 15N, 85°41’20W). ANSP 357487 A9753H T357.

Funa jeffreysi (Smith, 1875). South of Cape d’ Aguilar, Hong Kong, 20 m.

BM(NB).

Funa latisinuata (Smith, 1877). South of Cape d’ Aguilar, Hong Kong,

20 m. BM(NH).

Ptychobela suturalis (Gray, 1838). North Lantau Island, Hong Kong, 12 m.

BM(NH).

Cheungbeia mindanensis (Smith, 1877). South of Cape d’ Aguilar, Hong

Kong, 20 m. BM(NH).

Cheungbeia robusta (Hinds, 1839). South of Cape d’ Aguilar, Hong Kong,

20 m. BM(NH).

Vexitomina garrardi (Laseron, 1954). 2-3 km E. of Malabar, Sydney,

Australia (32°59.27’°S, 150°16.48’E). AM.

Inquisitor cf adenicus Sysoev, 1996, Off Ras Madrakah, Oman

(19°14.8°N, 56°26.8’E), 935 m. NMW.

Inquisitor aemula (Angas, 1877). Outer Lagoon, 10 m, Noumea, New

Caledonia. BM(NH).

Inquisitor latifasciata (Sowerby, 1870). Off Cape d’ Aguilar, Hong Kong,

25-30 m. BM(NH).

Epidirona gabensis (Hedley, 1922). 2 km East of Long Bay, Sydney, New

South Wales, Australia, 66 m. AM.

Antiguraleus morganus (Barnard, 1958). Off Mendu Point 300 m,

Transkei, South Africa (32°21.8’S, 29°0.0°E). NM.

Paraguraleus costatus (Hedley, 1922). 28 km east of south head of Little

Bay, Sydney, NSW, Australia (33°58°54"S, 151°33°38"E), 183-192 m.

Naudedrillia praetermissa (Smith, 1904). Off Sandy Point, Transkei,

South Africa (32°37.4’S, 28°36.9’E), 90 m. NM.

Nquma scalpta Kilburn, 1988. Off Park Rynie, Natal, South Africa

(30°23.2’S, 30°50.8’E), 140 m. NM.

Turridrupa bijubata (Reeve, 1843). 30 m, Beacon Island, Houtman

Abrolhos Islands, Western Australia. BM(NH).

Haedropleura septangularis (Montagu, 1803). Torre San Giovanni, BR,

Italy, 12 m. BM(NH).

Outgroup: subfamily Turrinae:

Gemmula deshayesii (Doumet, 1839). Off Cape d’ Aguilar, Hong Kong, 20

m. BM(NH).

Abbreviations: AM, Australian Museum; ANSP, Academy of Natural Sciences,

Philadelphia; BM(NH), Natural History Museum, London; NM, Natal Museum;

NMW, National Museum of Wales.

Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

are known from around the continental margins of the Indo-W.

Pacific Ocean (Wells, 1994; Taylor & Wells, 1994; Sysoev, 1996).

The relationships of the Crassispirinae to other conoideans are

uncertain and those of the genera within the Crassispirinae are

totally unknown; thus it is highly uncertain how the common genera

of the East Pacific such as Crassispira, Hindsiclava etc. are related

to some of the common Indo-W. Pacific genera such as Inquisitor,

and Funa.

In order to establish the anatomical range of the Crassispirinae

and evaluate characters which might be used to establish relation-

ships both within the group and with other conoideans, we studied

by serial sections the foreguts of 35 species. These were collected

within the East Pacific, Caribbean, Mediterranean and Indo-Pacific

provinces from a wide variety of habitats ranging from intertidal to

abyssal depths. Additionally, we used Gemmula deshayesi (sub-

family Turrinae) as an outgroup. This is the most extensive

comparative study yet attempted of the anatomy of any group of

Conoidea.

MATERIAL AND METHODS

Details of all the species used in the study are listed in Table 1. For

all species longitudinal serial sections were made of the foregut, cut

at 8 um and mostly stained in green Masson’s trichrome. Radulae

were cleaned with a dilute sodium hypochlorite solution and exam-

ined by SEM.

ANATOMICAL DESCRIPTIONS

Abbreviations used in the anatomical figures

asg acinous salivary gland

asgm modified acinous salivary gland

be buccal cavity

blp buccal lips

blpi _ invertible buccal lips

bm buccal mass

bs buccal sac

bt buccal tube

bts buccal tube sphincter,

btsa _ anterior sphincter of the buccal tube

btsi intermediate sphincter of the buccal tube

cf circular fold, surrounding opening of buccal sac

cfbt circular fold of the buccal tube

con _ circumoesophageal nerve ring

ct connective tissue of the buccal mass wall

cul connective tissue layer of the muscular bulb

ebt sac-ike enlargement of the buccal tube

ep epithelial pad

gre glandular part of the rhynchodeum

iep invaginated epithelium of the proboscis tip

ipt inverted proboscis tip

ire non-glandular invertible part of the rhynchodeum

Imb _ lumen of the muscular bulb

thickened muscular wall of sac-like enlargement of buccal

tube

ngre non-glandular non-invertible part of the rhynchodeum

oe oesophagus

oel oesophageal loop

p proboscis

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

pt proboscis tip

th/rhc rhynchocoel

rhs rhynchostomal sphincter

rs radular sac

rst rhynchostome

rtsg anastomosing tubular salivary gland

rw rhynchodeal wall

twg __ rhynchodeal wall glandular

sd salivary duct

se sac-like enlargement of buccal tube

sg salivary gland

stag single tube acinous salivary gland

t radular tooth.

tsg simple tubular salivary gland

Vv valvule

vg venom gland

vga duct of venom gland

In this section we describe and illustrate the foregut anatomy and

radulae of each of the species examined. The terminology of the

anatomical characters and organs largely follows Taylor ef al.,

(1993). Most of the features are illustrated diagrammatically in

Fig. 1, with details of others given in the descriptions of individual

species.

Crassispira (Crassispira) incrassata (Sowerby, 1834)

(Figs 2, 4a)

Rhynchodeum and proboscis

The rhynchostomal sphincter is large and located in a slightly

posterior position. The epithelium of the posterior rhynchodeal wall

btsa

ebt

btsi

Fig. 1 Composite diagram of the foregut of a hypothetical crassispirine

gastropod, showing the location of the major structures discussed in the

text. No single species has all these structures.

2y//

is continuous with that of the proboscis wall for slightly more than

half the length of the rhynchodeum. Extremely large proboscis

retractor muscles are attached to the rhynchodeum at the border

between the two different epithelia and at the proximal ends to the

columellar muscle.

The proboscis is long (more than half of the rhynchodeum),

slightly narrowing towards the tip. The proboscis tip is not inverted

inside. The muscles of the proboscis wall in the posterior two thirds

of the proboscis are significantly thicker than in the anterior section.

The proboscis lies coiled within the rhynchocoel (shown uncoiled in

drawing to illustrate relative proboscis length). There are two ante-

rior buccal tube sphincters. A sac-like enlargement of the buccal tube

is present, with the epithelium lining it taller than that of the rest of

the tube. An epithelial pad is also present. There is no intermediate

buccal tube sphincter.

The proboscis walls and posterior part of the rhynchodeum are

highly folded, suggesting significant elongation of the proboscis

during protraction. The proboscis wall is thick, comprising about

10% of proboscis diameter in the posterior half. In the anterior one

third of the proboscis, the wall is thinner, but due to the decrease in

proboscis diameter the wall comprises about 12% of the total

diameter. The wall of the buccal tube is also thick, comprising about

7% of proboscis diameter in its posterior half. Small buccal lips are

present.

Buccal mass and oesophagus

The large and very long buccal mass is located entirely within the

proboscis, with a thick wall and is curved. The oesophagus is

elongated between the buccal mass and nerve ring. The buccal sac is

very short.

Glands

The salivary glands are very large and acinous, protruding nearly to

the anterior of the rhynchodeum. The histology of the venom gland

changes abruptly before passing through the nerve ring. The duct of

the venom gland is ciliated, and opens into the buccal cavity at the

posterior border with the oesophagus. The muscular bulb is large,

with thick walls formed from two equal layers of circular muscle

fibres separated by a connective tissue layer, with a third much

thinner innermost layer of larger circular fibres.

Odontophore and radula

The odontophore is medium-sized, consisting of a pair of unfused

cartilages, formed by single layer of cells. The radula (Fig. 4a)

consists of marginal teeth of the robust wishbone type, with a thick,

distally-pointed major limb and a shorter, thinner minor limb. The

marginal tooth is short, ca.180um (0.5% of SL (shell length), 1.2%

AL (aperture length)).

Crassispira (Crassispira) maura (Sowerby, 1834)

(Figs 3, 4b)

Rhynchodeum and proboscis

The rhynchodeal sphincter is medium-sized and anteriorly located.

The epithelium of the anterior two thirds of the rhynchodeum is

glandular, but posteriorly, it changes abruptly to low, cubic non-

glandular, epithelium, which is continuous with that of the proboscis

wall. The anterior rhynchodeum is narrow with high folds.

The proboscis is short, occupying less than half the length of the

rhynchodeum, with the proboscis tip not inverted. The proboscis

walls are rather thick, forming about 20% of the proboscis diameter,

but the walls of the buccal tube are thin, composing about 4% of the

total diameter. The mouth is narrow. The muscles of the proboscis

58 Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

, — — —

(= nN \ Ss

f, S

s &

Fig. 2 Crassispira (Crassispira) incrassata (Sowerby, 1834). A, semidiagrammatic longitudinal section of the foregut (only one salivary duct is shown);

B, longitudinal section of the proboscis tip.

Fig. 3 Crassispira (Crassispira) maura (Sowerby, 1934). A, semidiagrammatic longitudinal section of the foregut (salivary glands and ducts not shown);

B, longitudinal section of the proboscis and buccal mass.

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

o —

Se)

Fig. 4 Radulae of Crassispirinae. a, Crassispira (Crassispira) incrassata b, Crassispira (Crassispira) maura c, Crassispira (Gibbaspira) dysoni d, C.

(Glossispira) harfordiana flucki. Scale bars = 20um.

walls are equally developed along its length. There is a small anterior

buccal tube sphincter, positioned at the base of the small sac-like

enlargement of the buccal tube, which also has an epithelial pad. An

intermediate buccal tube sphincter is absent. The buccal tube is very

narrow, both inside the proboscis and for some distance behind it,

and lined with a very low epithelium. Some distance behind the

proboscis, the buccal tube expands greatly and forms two small,

poorly-muscularized lips which are directed anteriorly, similar to

the ‘valvule.’ (Sheridan et al. 1973). The buccal lips are large and

muscular, with the dorsal one inverted inside the buccal cavity.

Buccal mass and oesophagus

The buccal mass is medium-sized and situated to the posterior of the

proboscis base. Its dorsal wall is very thin. By contrast, the ventral

wall near the entrance of the radular diverticulum is thick, but

formed mainly by a layer of loose connective tissue, which is 4-8

times thicker than the muscle layer. The oesophagus is greatly

elongated between the buccal mass and nerve ring, forming a long

loop. The buccal sac is of medium length and narrow.

Glands

The salivary glands are very large and acinous, with ducts that are

thick and coiled, but become very thin as they approach the buccal

mass. The venom gland changes in histology after passing anteriorly

through the nerve ring. The duct of the gland is narrow, unciliated,

and opens just to the posterior of the buccal cavity. The muscular

bulb is large, with most of the wall formed by two subequal layers of

longitudinal muscle fibres, divided by a connective tissue layer, with

a third innermost, thin layer of circular muscle fibres.

Odontophore and radula

The odontophore is medium-sized with paired, unfused cartilages.

The radula consists of only marginal teeth of the wishbone type, with

the major limb robust, solid and pointed with a long straight leading

edge. The minor limb is smaller and thinner, but tapers towards the

base (Fig. 4b). The marginal tooth is short, ca.130um (0.3% of SL,

1.0% AL).

Crassispira (Gibbaspira) dysoni (Reeve, 1846)

(Figs 4c, 5)

Rhynchodeum and proboscis

The rhynchostomal sphincter is medium-sized and anteriorly lo-

cated. The epithelium of the anterior half of the rhynchodeal wall is

tall and glandular, while the posterior half is continuous with that of

the proboscis wall. The proboscis tip is not inverted. The proboscis

is short (about half of the rhynchocoel), not coiled, but slightly

folded and narrowing towards the tip. The muscles of the proboscis

wall are equally developed along its length.

The anterior buccal tube sphincter is small, whilst the sac-like

enlargement of the buccal tube is long, with a long epithelial pad,

formed of large cubic cells with large nuclei. At the base of the

enlargement, there is an intermediate sphincter, which lies at a

distance of three radular tooth lengths from the mouth. The walls

of the sac-like enlargement are thicker than the rest of the buccal

tube and similar to those of C. (Crassiclava) spp. The buccal tube

is lined with ciliated epithelium. The proboscis wall is thin, form-

ing about 10% of the proboscis diameter, while the wall of the

buccal tube comprises about 7% of proboscis diameter. Buccal

lips are absent.

Buccal mass and oesophagus

The buccal mass lies to the posterior of the proboscis, and is small,

thin-walled and curved, and comprises less than one fifth of the

gre

Nits >

Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

proboscis length. The oesophagus is greatly elongated between the

buccal mass and nerve ring and forms a long loop. The epithelium

bears very long cilia, which occupy nearly the whole lumen. The

opening of the radular diverticulum into the buccal cavity is quite

narrow and bordered by a rather tall circular muscular fold, similar

to that seen in C. harfordiana. The salivary ducts open into the

radular sac at the base of this fold (Fig. 5).

Glands

The salivary glands are large and acinous, with ducts that are

thick, very long and highly coiled. The histology of the venom

gland changes after passing anteriorly through the nerve ring. The

duct of the gland is narrow, highly coiled and probably ciliated.

The gland itself is very long, thick, and occupies a large part of the

body haemocoel. The muscular bulb is large, with thick walls

formed of two equal layers of circular muscle fibres, divided by a

connective tissue layer. The lumen of the bulb is filled with venom

granules.

Odontophore and radula

The odontophore is small, consisting of paired, unfused, subradular

cartilages, formed by a single layer of cells. The radula consists of

marginal teeth (Fig. 4c) which are of the wishbone type, with a large,

solid, sharply pointed, major limb and a shorter, slender, secondary

limb. The marginal tooth is medium long, ca.105um (0.8% of SL,

1.8% AL).

Mivsthai ls letidyy zy)

Be aR

61; ou

Cy ela

adage AAA ecb

Fig.5 Crassispira (Gibbaspira) dysoni (Reeve, 1846). A, semidiagrammatic longitudinal section of the foregut (salivary ducts not shown); B, longitudinal

section of the proboscis tip; C, section of the buccal mass showing the opening of the radular sac.

——— ——adll

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

“

AO Doers

Fig.6 Crassispira (Glossispira) harfordiana flucki (Brown & Pilsbry, 1913). A, semidiagrammatic longitudinal section of the foregut (salivary glands not

shown); B, longitudinal section of the proboscis tip; C, section of the buccal mass, showing the opening of the radular sac.

Crassispira (Glossispira) harfordiana flucki (Brown &

Pilsbry, 1913)

(Figs 4d, 6, 30e)

Rhynchodeum and proboscis

The rhynchostomal sphincter is small and slightly posteriorly situ-

ated. The epithelium of the rhynchodeum wall is glandular and that

of the posterior rhynchodeal wall is not continuous with that of the

proboscis wall. The proboscis tip is not inverted. The proboscis is

very long and highly coiled (longer than the rhynchocoel and shown

uncoiled in Fig. 6). The muscles of the proboscis wall are equally

developed along its length, except at the very anterior.

The anterior part of the proboscis from the mouth opening to the

buccal mass is very thin (about 0.1 mm compared to the proboscis

length of about 5—6 mm) and highly folded. The proboscis tip is

highly expanded to form a wineglass-shaped structure (Fig. 6, 30e).

Closer to the tip, there is a septum with a small, circular sphincter

surrounding the narrow opening. The septum delimits the sac-like

enlargement of the buccal tube. The epithelium lining this is signifi-

cantly taller than that of the rest of the tube, forming a low epithelial

pad. At the base of the enlargement, there is a second small sphincter.

Pieces of one, or possibly more, marginal teeth (fragmented during

sectioning) were seen attached to the epithelium. The distance

between the two sphincters corresponds to about one tooth length. It

is possible, that the enlargement serves as a storage area for detached

marginal teeth. There is no intermediate sphincter in the buccal tube.

Buccal cavity and oesophagus

The buccal mass is long (equivalent to about one quarter of probos-

cis length), with moderately thick walls and no curvature. It is

contained entirely within the proboscis and starts within the distal

third of the proboscis length. The walls of the buccal cavity form

several annular folds. The buccal lips are small.

The oesophagus is highly elongated between the buccal mass and

nerve ring, but because the buccal mass is located anteriorly within

the proboscis, there is virtually no loop. A buccal sac is absent. The

opening of the radular diverticulum into the buccal cavity is narrow

and bordered by a rather tall circular muscular fold. The salivary

ducts open into the cavity at the base of this fold (Fig. 6).

Glands

The salivary glands are small, paired and acinous and lie at the

proboscis base. The ducts are highly coiled. The histology of the

venom gland changes abruptly after passing through the nerve ring.

The duct of the venom gland is unciliated, narrow, and highly coiled

before opening into the buccal cavity at the border with the oesopha-

gus. The muscular bulb is large, with thick walls formed of two

layers of longitudinal muscle fibres, separated by a connective tissue

layer, with a much thinner innermost layer of larger circular fibres

Odontophore and radula

The odontophore is small and composed of a pair of unfused

cartilages formed by a single layer of cells. The radula consists of

marginal teeth of the wishbone type (Fig. 4d), with a solid, pointed,

curved major element and a shorter and narrower secondary ele-

ment. The marginal tooth is medium long, ca.195um (0.7% of SL,

2.7% AL).

Crassispira (Crassispirella) latizonata (E. A. Smith, 1882)

(Fig. 7)

Sp)

elon ee

Va?

Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

Fig. 7 Crassispira (Crassispirella) latizonata (Smith, 1882). Longitudinal section of the posterior part of the proboscis and buccal mass (only one salivary

duct is shown).

Rhynchodeum and proboscis

The rhynchostomal sphincter is small and anteriorly located. The

epithelium of the anterior half of the rhynchodeal wall is glandular,

arranged into high folds, but to the posterior it is continuous with that

of the proboscis wall. The dorsal wall of the proboscis tip is inverted

inside. The proboscis is medium long (about half of the rhynchocoel)

and not coiled. The muscles of the proboscis wall are equally

developed along its length.

There is a small, anterior buccal tube sphincter, which lies in front

of the well developed, sac-like enlargement of the buccal tube, lined

with tall epithelium. There is no epithelial pad. A large, intermediate

sphincter of the buccal tube is present, which lies at the base of the

distal enlargement at a distance of about 2.5 radular tooth lengths

from the mouth opening. The proboscis walls are not thick, compos-

ing about 15% of proboscis diameter. The wall of the buccal tube

wall is also not thick, forming about 8% of proboscis diameter.

Small buccal lips are present.

Buccal cavity and oesophagus

The buccal mass lies posterior to the proboscis base and is large,

equivalent to about one quarter of proboscis length. Anterior to the

opening of the radular sac, the walls of the buccal cavity are thin,

similar to those of the buccal tube, but become thicker to the

posterior.

The oesophagus is greatly elongated between the buccal mass and

nerve ring and forms a very long loop. A remarkable feature of the

oesophagus 1s that its epithelium bears very long cilia which occupy

the whole lumen. The duct of the radular sac is broad and rather long.

The buccal sac very short.

Glands

The salivary glands are tubular in histology, coiled and situated near

the nerve ring. The salivary ducts are very long, coiled and thick. The

histology of the venom gland changes abruptly after passing through

the nerve ring. The duct of the venom gland is ciliated, coiled and

narrow, with the same diameter as the salivary ducts, and opens into

the buccal cavity at the border with the oesophagus. The muscular

bulb is large, with thick walls formed of two layers of longitudinal

muscle fibres, divided by a connective tissue layer, with a much

thinner, innermost layer of circular fibres.

Odontophore and radula

The odontophore is rather large and consists of paired subradular

cartilages, formed by single layer of cells. The radula was not

examined and there are, unfortunately, no published illustrations.

Crassispira (Monilispira) pluto Pilsbry and Lowe, 1932

(Fig. lla )

Rhynchodeum and proboscis

The rhynchostomal sphincter is large and anteriorly situated. The

epithelium of the anterior seven eighths of the rhynchodeal wall is

tall, glandular and arranged into high folds, while that of the

posterior one eighth is continuous with the proboscis wall. The

proboscis tip is not infolded. The proboscis is very long (ca 1.5 times

longer than rhynchocoel), coiled in the anterior part and thick. The

muscles of the proboscis wall are more developed at the base.

An anterior buccal tube sphincter is present. The sac-like enlarge-

ment of the buccal tube is present, but not well differentiated. Also,

there is an epithelial pad with a marginal tooth attached to it. An

intermediate sphincter is absent. The proboscis walls are thick,

comprising about 20% of proboscis diameter, whilst the buccal tube

wall is medium-thick, forming about 15% of the total diameter. The

buccal lips are large and muscular.

Buccal mass and oesophagus

The buccal mass lies posterior to the proboscis, is rather long, with

thick walls, uncurved and equivalent to about half the proboscis

length. The oesophagus is elongated between the buccal mass and

nerve ring and forms a long loop. The buccal sac is very short.

Glands

The salivary glands are large and acinous. The histology of the

venom gland changes after passing the nerve ring. The duct of the

venom gland is narrow and unciliated. The gland itself is long. The

muscular bulb was unfortunately missing from the sections.

Odontophore and radula

The odontophore is large, consisting of paired, unfused, subradular

cartilages, formed by a single layer of cells. The radula consists of

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

Fig. 8 Crassispira (Striospira) tepocana Dall, 1919. Semidiagrammatic longitudinal section of the foregut (salivary ducts and glands not shown).

marginal teeth of the wishbone type (Fig. | la), with a solid, sharply-

pointed major limb and a shorter, slender, secondary limb. The

marginal tooth is long, ca.175um (1.1% of SL, 2.9% AL).

Crassispira (Striospira) tepocana Dall, 1919

(Fig. 8)

Rhynchodeum and proboscis

The rhynchostomal sphincter is small and anteriorly located. The

epithelium of the anterior two thirds of the rhynchodeal wall is tall

and glandular, while that of the posterior one third is continuous with

the proboscis wall.

The proboscis is short (about half of the rhynchocoel), coiled and

folded in its anterior part, becoming narrow towards the tip. The

muscles of the proboscis wall are better developed at the base. The

proboscis tip is not infolded. There is no anterior buccal tube

sphincter and the sac-like enlargement of the buccal tube is slight.

The anterior part of the buccal tube is very narrow. There is a very

small intermediate sphincter, which lies at a distance of about three

radular tooth lengths from the very narrow mouth opening. The

proboscis wall is thick, composing 25% of proboscis diameter at its

base, and about 16% in its apical part. The buccal tube wall is

medium-thick, making up about 10% of proboscis diameter. The

buccal lips are very small.

Buccal mass and oesophagus

The buccal mass is long and curved and lies posterior to the

proboscis, equivalent to about two thirds of proboscis length. The

oesophagus is elongated between the buccal mass and nerve ring and

forms a short loop. The epithelium bears very long cilia, which

occupy nearly the whole lumen. The duct of the radular sac is broad

and very short, whilst the buccal sac is virtually absent.

Glands

The salivary glands are medium-sized, with the ramified tubular

morphology. The histology of the venom gland changes abruptly

anterior to the nerve ring. The duct of the venom gland is narrow and

unciliated. The gland itself is very long and occupies a large part of

the body haemocoel. The muscular bulb is extremely large, with

thick walls formed of two subequal layers of longitudinal muscle

fibres, divided by a connective tissue layer, with a much thinner

innermost layer of circular muscle.

Odontophore and radula

The odontophore is medium-sized, with paired, unfused subradular

cartilages, formed by a single layer of cells. The radula is illustrated

by McLean (1971, fig. 66) and consists of marginal teeth of the

wishbone type (similar to C. kluthi, Fig. 11b), with a large, robust

and pointed major limb and a thinner, secondary element which is

attached near the tip of the major limb.

Crassispira (Striospira) kluthi Jordan, 1936

(Fig. 11b)

The foregut anatomy of this species is similar to that of C. tepocana

and only the differences are listed below.

The rhynchostomal sphincter is larger and located further to the

posterior. The rhynchodeum is entirely glandular with no change in

epithelium along its length. The proboscis is highly coiled. The

proboscis walls are very thin in the anterior part and are thinner than

in C. tepocana (about 15 % of the proboscis diameter). The buccal

mass lies within the proboscis base. The oesophagus is more greatly

elongated and coiled between the buccal mass and nerve ring. The

duct of the venom gland is very long and highly coiled. The radular

teeth (Fig. 11b) have a large, pointed, major limb and a more slender

secondary limb which is attached near the tip of the major limb. The

marginal teeth are short, ca.90um (0.5% of SL, 1.4% AL).

Crassispira (Striospira) xanti Hertlein & Strong, 1951

(Fig. 9)

Rhynchodeum and proboscis

The rhynchodeal sphincter is large and anteriorly located. The

epithelium of the anterior half of the rhynchodeum is glandular and

arranged into high folds; posteriorly, this changes abruptly to a non-

glandular epithelium, which is continuous with that of the proboscis

wall.

The proboscis is short, about half of the rhynchodeum length,

with the tip not infolded. The proboscis walls are thick, composing

64 Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

btsi cfbt

Fig.9 Crassispira (Striospira) xanti Hertlein & Strong, 1951. A, longitudinal section of the proboscis and buccal mass (salivary ducts not shown): B,

longitudinal section of the proboscis tip.

Fig. 10 Crassispira (Crassiclava) turricula (Sowerby, 1834). Semidiagrammatic section of the foregut.

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

about 25% of proboscis diameter, whilst the buccal tube walls are

highly folded, but thin, making up about 6% of the total diameter.

The mouth is narrow. The proboscis is highly folded, with the base

more muscular than the tip. Both the anterior buccal tube sphincter

and the epithelial pad are absent. The sac-like enlargement of the

buccal tube is long and lined with tall epithelium, but poorly

differentiated from the buccal tube. It has two unequal circular folds.

The anterior one is larger and directed anteriorly and lined with tall

epithelium. To the posterior of the second fold, the wall of the buccal

tube forms an invagination, where the base of the radular tooth is

situated. The intermediate sphincter of the buccal tube is rather

large. The buccal tube is lined with a low, loose epithelium and after

leaving the proboscis it expands slightly in diameter and forms two

small, poorly-muscularized lips which are directed anteriorly, simi-

lar to the ‘valvule’ seen in Mangeliinae (Sheridan ef al., 1973). The

buccal lips are medium-sized, muscular, with the dorsal one inverted

inside the buccal cavity.

Buccal mass and oesophagus

The buccal mass is situated to the posterior of the proboscis, is rather

large and muscular, with a narrow lumen, and sharply curved. The

oesophagus is greatly elongated between the buccal mass and nerve

ring, forming a long loop. The buccal sac is long and narrow.

Glands

The salivary glands are medium-sized and acinous, with long, coiled

and rather thick ducts. The venom gland changes in histology after

passing through the nerve ring, but its diameter remains nearly the

same and there is no defined duct. The gland opens into the oesopha-

gus just posterior to the buccal cavity. The muscular bulb is large,

composed of two layers of longitudinal muscle fibres, divided by a

connective tissue layer, with an innermost, thin layer of circular

muscle fibres.

Odontophore and radula

The odontophore is medium-sized, with paired unfused cartilages

formed of one layer of cells. The radula illustrated by McLean

(1971, fig. 67) is composed of marginal teeth of the wishbone type,

with a robust and pointed major limb and a smaller, shorter, second-

ary limb.

Crassispira (Crassiclava) turricula (Sowerby, 1834)

(Figs 10, 11c—d)

Rhynchodeum and proboscis

The rhynchostomal sphincter is large and anteriorly situated. The

epithelium of the anterior one third of the rhynchodeal wall is

glandular, and arranged into high folds; whilst the posterior two

thirds is continuous with that of the proboscis wall. The proboscis tip

is formed by the invagination of the outer proboscis wall into the

buccal tube. This is confirmed by the similar, low, cubic epithelia of

the outside of the proboscis and the anterior-most part of the buccal

tube. Posteriorly in the buccal tube the cubic epithelium is replaced

by one with columnar cells.

The proboscis is of medium length (little more than half of the

thynchocoel), uncoiled, but with telescopically folded walls. The

muscles of the proboscis wall are better developed at its base.

An anterior buccal tube sphincter is present. The sac-like enlarge-

ment of the buccal tube is well-defined and lined with tall, glandular

epithelium. An intermediate sphincter is also present, situated at the

base of the sac-like enlargement. The walls of the buccal tube in the

area of the enlargement are much thicker and formed by relatively

thicker circular muscle fibres, as if this part of the tube is capable of

strong contraction. The proboscis wall is thick, composing about

25% of proboscis diameter, whilst the wall of the buccal tube is

medium-thick, representing about 5% of proboscis diameter. Large

muscular buccal lips are present.

Buccal mass and oesophagus

The buccal mass lies mainly within the proboscis and is very long,

equivalent to about two thirds of the proboscis length. The oesopha-

gus is greatly elongated between the buccal mass and nerve ring and

forms a long loop. It is narrow and lined with an epithelium bearing

long cilia. The duct of the radular sac is broad and rather long. The

buccal sac is short.

Glands

The salivary glands are large, with the ramified tubular morphology.

The salivary ducts are very long, coiled, and thick. The histology of

the venom gland changes anterior to the nerve ring. The duct of the

venom gland is narrow and unciliated. The muscular bulb is ex-

tremely large, with thick walls mainly formed of two layers of

longitudinal muscle fibres, separated by a connective tissue layer,

with a much thinner, innermost layer of circular fibres.

Odontophore and radula

The odontophore is large, consisting of paired, unfused, subradular

cartilages, formed by a single layer of cells. The radula (Figs 1 1c—d)

consists of both lateral and marginal teeth. The lateral teeth ( use of

the term ‘lateral’ teeth here does not imply homology with the lateral

teeth of other neogastropods) are low and arcuate, sharply-curved

anteriorly towards the midline of the ribbon. In profile these teeth are

cuesta-like, with steep anterior faces and gentle posterior slopes.

The marginal teeth are of the robust wishbone form with a large,

pointed, major element and a thinner, shorter, minor element. The

marginal teeth are ca 200um long (0.6 %SL; 2.0% AL)

Crassispira (Crassiclava) apicata (Reeve, 1845)

(Fig. 12)

The anatomy of the foregut is similar to the preceding species and

only the differences are mentioned.

The anterior part of the rhynchodeum, which is lined with glandu-

lar epithelium is shorter than in C. turricula and comprises about

half the length of the rhynchodeum. The proboscis walls do not form

telescopic folds. Also, the proboscis is longer and occupies about

two thirds of the rhynchocoel. Only the anterior buccal tube sphinc-

ter is present and an epithelial pad is located within the sac-like

enlargement of the buccal tube. The enlargement itself is shorter

than in C. turricula. The proboscis walls are much thinner and

comprise only about 6% of its diameter. The salivary glands are very

large with the ramified tubular morphology. The radula (from Maes,

1983 figs 31 & 37) consists of both lateral and marginal teeth. The

lateral teeth have the arcuate form illustrated for C. turricula (Figs

11c—d), whilst the marginal teeth have the wishbone form with a

robust major limb and a shorter, thinner, minor limb.

Hindsiclava andromeda (Dall 1919)

(Fig. 13)

Rhynchodeum and proboscis

The rhynchostomal sphincter is of medium size and located slightly

to the posterior. The epithelium of the anterior half of the rhynchodeal

wall is tall and glandular, whilst the posterior is low and cubic and

Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

Fig. 11 Radulae of Crassispirinae. a, Crassispira (Monilispira) pluto, scale bar = 20um b, Crassispira (Striospira) kluthi scale bar = 1Oum c, Crassispira

(Crassiclava) turricula scale bar = 30 um d, C. (C.) turricula scale bar = 20 um.

continuous with that of the proboscis wall. The wall of the

rhynchodeum in its posterior part is muscular and free and much

thicker than to the anterior. This posterior part of the rhynchodeum

is able to evert. Powerful retractor muscles are attached at the point

where the rhynchodeal epithelium changes; these run along the

rhynchodeum and are attached to the buccal mass.

In the retracted position, the proboscis is rudimentary (Fig. 13),

being only about twice as long as than the radular tooth length. The

mouth opening is very narrow. There is no anterior buccal tube

sphincter and the sac-like enlargement of the buccal tube is small. At

the base of the enlargement there is an intermediate sphincter, which

lies at a distance of about three radular tooth lengths from the mouth

and posterior to the proboscis. The buccal tube forms a loop poste-

rior to the proboscis and widens greatly before opening in the buccal

mass.

Buccal mass and oesophagus

The buccal mass is medium-sized, and located to the posterior of the

Fig. 12

bisie!

Crassispira (Crassiclava) apicata (Reeve, 1845).

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

ESI

GED?

Fig. 13. Hindsiclava andromeda (Dall, 1919). A, Semidiagrammatic longitudinal section of the foregut (salivary ducts not shown); B, longitudinal section

of the proboscis and buccal mass (only one salivary duct is shown).

proboscis. It has rather muscular walls and is not curved. The buccal

lips are asymmetrical, the ventral lip being large, and the dorsal

medium-sized. The oesophagus is greatly elongated between the

buccal mass and nerve ring and forms a long loop. The opening of

the radular diverticulum into the buccal cavity is narrow, and the

salivary ducts open at the entrance of radular diverticulum into the

buccal mass. A buccal sac is absent.

Glands

The salivary glands are very large with the ramified tubular mor-

phology. The salivary ducts are very long and thick, but only slightly

coiled. The histology of the venom gland changes abruptly soon

after passing anteriorly through the nerve ring. The duct of the

venom gland is narrow, coiled and unciliated. The muscular bulb is

large, with a narrow lumen and thick walls mainly formed of two

equal layers of longitudinal muscle fibres, divided by a connective

tissue layer, with a thin, innermost layer of circular fibres.

Odontophore and radula

The odontophore is small, consisting of a pair of unfused, subradular

cartilages, formed by a single layer of cells. Unfortunately, no

radular information is available for this species, but it is presumed to

be similar to Hindsiclava militaris (see below).

Hindsiclava militaris (Reeve, 1843)

(Figs 14 a—b)

Two specimens were sectioned. The species is very similar in

general foregut anatomy to H. andromeda and differs mainly in the

slightly longer proboscis (which occupies from 1/7 to 1/5 of the

rhynchodeum). The proboscis is very muscular, with the entire

lumen filled with retractor muscles. The anterior buccal tube sphinc-

ter is very small, and hardly visible. The proboscis tip is invaginated.

This is confirmed by the change of the epithelium in the anterior part

of the buccal tube. A short buccal sac is present.

Radula

The radula consists mainly of two rows of marginal teeth. These are

wishbone in form but have a very distinctive, hairpin-like morphol-

ogy (Figs 14 a—b). The major limb is elongate and pointed, slightly

concave-upwards in profile with raised edges. The secondary limb is

long and slender, attached to the major limb near the tip but detached

and distantly separated for most of the length, with separate attach-

ment to the radular membrane. Another feature of the radula is the

regular low, transverse ridges which cross the central part of the

ribbon (Fig. 14a) The marginal teeth are medium long, ca.140um

(0.5% of SL, 1.9% AL).

Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

Fig. 14 Radulae of Crassispirinae a, Hindsiclava militaris b, H. militaris c, Miraclathurella bicanalifera d, Haedropleura septangularis. Scale bars =

20um.

Miraclathurella bicanalifera (Sowerby, 1834)

(Fig. 14c)

Rhynchodeum and proboscis

The rhynchodeal sphincter is small and anteriorly located. The

epithelium of the anterior part of the rhynchodeal cavity is glandular,

whilst that of the posterior rhynchodeum is low, non-glandular, and

continuous with that of the proboscis wall The rhynchostome is

rather wide.

The proboscis is as long as the rhynchodeal cavity, highly folded,

and very thick at the base, but sharply narrowing towards the tip. The

proboscis walls are thick and form about 22% of the proboscis

diameter at its base. The wall muscles at the proboscis tip are much

thinner. The proboscis is very muscular and its lumen is mostly filled

with the retractors. The mouth is narrow. The anterior buccal tube

sphincter is absent. The sac-like enlargement of the buccal tube is

poorly defined, but lined with taller epithelium with a small epithe-

lial pad. There is a large intermediate sphincter, situated approxi-

mately one third of the distance down the proboscis. The buccal tube

is very narrow anteriorly, but broad in the posterior part of the

proboscis. Its walls are highly folded and compose about 10% of the

proboscis diameter. The buccal tube is lined with a very low epithe-

lium, which is replaced with a tall one, continuous with that of the

buccal cavity, slightly anterior to the buccal cavity.

Buccal mass and oesophagus

The buccal mass is large, slightly less that half of the proboscis

length, and lies within the proboscis. It has rather thick walls with no

curvature. The buccal lips are very small and poorly defined. Ante-

rior to them, the walls of the buccal tube form a fold, similar in

appearance to the true buccal lips. The oesophagus is elongated

between the buccal mass and nerve ring and forms a medium-long

loop. A buccal sac is absent. The salivary ducts open in the buccal

cavity on both sides of the very broad opening of the radular

diverticulum.

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

Glands

The salivary glands are large, paired and acinous. The venom gland

changes in histology after passing anteriorly through the nerve ring.

The duct is very narrow, ciliated, coiled, and opens at the border

between the buccal mass and oesophagus. The muscular bulb is

large, with the wall formed of two equal layers of longitudinal

muscle fibres, divided by a connective tissue layer, with an inner-

most, thin layer of circular muscle.

Odontophore and radula

The odontophore is rather large and protrudes into the buccal cavity.

It has paired, unfused cartilages, formed by single layer of cells. The

radula (Fig. 14c) consists of marginal teeth of the wishbone type,

each tooth with a large major limb with a pointed tip, the middle part

of the tooth broad and concavo/convex and narrowing towards the

base. The secondary limb is thin at the distal end broadening towards

the base. The marginal tooth is 172um in length (1.1% of SL, 4.2%

AL)

Haedropleura septangularis (Montagu, 1803)

(Figs 14d, 15)

The specimen sectioned was rather similar to the illustration in

Sheridan et al. (1973, fig. 6) and differs only in some details.

Rhynchodeum and proboscis

The rhynchodeal sphincter is small and anteriorly located. The

epithelium of the anterior half of rhynchodeum wall is tall, glandular

and folded. Posteriorly, it is replaced abruptly with a low, non-

glandular, cubic epithelium continuous with that of the proboscis

wall.

The whole rhynchocoel is rather short and the thick proboscis,

although not long, occupies nearly the entire cavity. The ratio

between the proboscis length and its diameter is about 1.3. The

proboscis tapers toward the tip, which is slightly invaginated at the

mouth opening. The proboscis walls are medium-thick and compose

about 16% of the proboscis diameter at its base. The mouth opening

is very narrow. The muscles of the proboscis wall are equally

developed along the length, and only in the inverted part are they

somewhat thinner. The proboscis retractor muscles are very large

and occupy the whole inner lumen. A large anterior buccal tube

sphincter is present, situated somewhat posterior to the proboscis

tip. The distance between the sphincter and the uninverted part of the

proboscis tip is equivalent to the length of a single radular tooth. A

sac-like enlargement of the distal part of the buccal tube is present,

but poorly defined. This is lined with loose, tall, ciliated epithelium,

while the remaining part of the tube is lined with extremely low,

Fig. 15 Haedropleura septangularis (Montagu, 1803). Longitudinal

section of the proboscis (radular sac not shown).

inconspicuous, epithelial cells. There is no intermediate sphincter.

The buccal tube has thin walls.

Buccal mass and oesophagus

The buccal mass lies to the posterior of the proboscis base, and is

rather long, equivalent to about two thirds of the proboscis length. It

has thick walls and a narrow inner cavity, which is uncurved. Long,

extensible, buccal lips are present (not shown by Sheridan et al.,

1973). The oesophagus shows no elongation between the buccal

mass and nerve ring.

Glands

The salivary glands are paired, large, tubular and coiled. Their

diameter is much larger than illustrated by Sheridan et al. (1973) and

only 2—3 times smaller than that of the venom gland. The salivary

ducts are very short, narrower than the glands and ciliated. There is

no change in the histology of the venom gland to the anterior of the

nerve ring, and it opens into posterior part of the buccal cavity at the

boundary with the oesophagus. The venom gland is very large,

highly coiled and occupies most of the body haemocoel. The muscu-

lar bulb is very large and elongate; longer and thicker in fact than the

proboscis. Its wall is formed of two subequal layers of longitudinal

muscle fibres, divided by a connective tissue layer, with a very thin

innermost layer of circular fibres.

Odontophore and radula

The odontophore is medium-sized, with paired, unfused,

odontophoral cartilages. The radula consists of marginal teeth,

which are of the wishbone type (Fig. 14d), but with the major

element having a large, spathulate, pointed, distal end and a narrow

shaft. The minor element is thinner and attaches distally to the broad

blade of the major element and to the radula ribbon at the base. The

marginal tooth is long, ca.145um (1.7% of SL, 4.0% AL).

Nquma scalpta Kilburn, 1988

(Figs 16, 17a, 30a)

Rhynchodeum and proboscis

The rhynchodeal sphincter is large and anteriorly situated. The

epithelium of the anterior two thirds of the ventral rhynchodeum

wall and three quarters of the dorsal wall is tall, glandular, folded and

formed of large cells. The epithelium of the posterior rhynchodeum

is formed of low, non-glandular, cubic cells continuous with that of

the proboscis wall.

The proboscis is long, slightly longer than the rhynchocoel and

with the tip not infolded. The proboscis walls are medium-thick,

comprising about 13% of the proboscis diameter in its central part.

The mouth opening is extremely narrow. The muscles of the probos-

cis wall are equally developed along its posterior part, but are absent

near the distal tip of the proboscis. There is no anterior sphincter of

the buccal tube. The distal end of the buccal tube possesses a sac-like

enlargement which is lined in the anterior part with very tall and

narrow columnar cells where there is also an epithelial pad. The base

of a marginal radular tooth was seen attached to the pad. Posterior to

the pad, the epithelium is very low, similar to that of the remaining

part of the buccal tube, but becomes tall and probably glandular just

in front of the large intermediate sphincter, which lies close, about

two marginal tooth lengths, distant from the proboscis tip. The

buccal tube has rather thin walls.

Buccal mass and oesophagus

The buccal mass is long, equivalent to about 2/3 of the proboscis

length and lies at the proboscis base, projecting a long way to the

ire bm

ee aA

LUT — 7

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———— SS N\

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Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

Fig. 16 Nquma scalpta Kilburn, 1988. A, Semidiagrammatic longitudinal section of the foregut (salivary ducts not shown); B, longitudinal section of the

proboscis tip showing a gripped marginal tooth.

posterior of the rear of the proboscis. It has relatively thin walls and

a narrow inner cavity which is uncurved. Extensible buccal lips are

absent. The oesophagus is elongated between the buccal mass and

nerve ring, forming a rather long loop.

Glands

The salivary glands are large, fused and acinous. The ducts are

paired, very long, coiled, thick and leave the gland at the same place.

The venom gland changes histology after its anterior passage through

the nerve ring. The duct of the gland is narrow, ciliated, long and

coiled, and opens into posterior part of the buccal cavity. The wall of

the muscular bulb is formed of three layers, the two outermost being

divided by a connective tissue layer, with a very thin innermost layer

of circular fibres. The two outer layers are formed from longitudinal

muscle fibres, the outer being twice as thick as the inner.

Odontophore and radula

The odontophore is small, with paired and unfused, odontophoral

cartilages. The buccal sac is short. The radula (Fig. 17a) consist of

marginal teeth, with each wishbone tooth formed of two elements.

The major element is robust and solid with a distal point. The

secondary element is smaller (0.6 of the major element), thinner and

terminates before the base of the tooth. The marginal tooth is

medium long, ca.130um (1.3% of SL, 2.7% AL).

Naudedrillia praetermissa (Smith, 1904)

The foregut anatomy is similar in general pattern to that of Nqguma

scalpta and differs in the position of the buccal mass, which lies

totally posterior to the proboscis, and the more developed but

nevertheless still short buccal lips. The striking difference is that in

N. praetermissa the salivary glands are tubular, large and highly

coiled, with a rather wide inner lumen. The ducts are very short, of

similar diameter to the glands, but ciliated.

A radular tooth was seen in the sac-like enlargement of the buccal

tube, attached to the epithelial pad. The radula consists of marginal

teeth (Kilburn, 1988, fig. 52), with each tooth comprising two

components; a large major element which is an elongate, grooved

and twisted blade, with a distal point and a shorter minor element

which is attached about halfway along the major element. The

marginal tooth is long, ca.160um (1.3% of SL, 3.5% AL).

Epidirona gabensis (Hedley, 1922)

(Fig. 18a)

Rhynchodeum and proboscis

The rhynchostomal sphincter is large and posteriorly located. The

rhynchocoel is very long and narrow. The epithelium of nearly the

whole rhynchodeal cavity is tall, glandular, folded and formed of

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

vil

Fig. 17 Radulae of Crassispirinae. a, Nquma scalpta scale bar = 20um b, Inquisitor latifasciata scale bar = 25 um c, Inquisitor aemula scale bar = 20 um

d, Inquisitor aff. adenicus scale bar = 20 um.

large cells. In approximately the posterior one tenth of the

rhynchodeum the epithelium is non-glandular and continuous with

that of the proboscis wall.

The proboscis is short in comparison with the rhynchocoel (about

1/5 of its length), rather thick and not coiled. The proboscis walls are

thick, forming about 25% of the proboscis diameter. The mouth is

very narrow. The muscle of the proboscis walls are equally devel-

oped along its length. The anterior buccal tube sphincter is very

small and lies posterior to the mouth in front of medium-sized sac-

like enlargement of the buccal tube. The epithelium, lining the

enlargement forms a large pad, to which the base of a marginal tooth

was attached. There is a large intermediate sphincter, situated at

about three marginal tooth lengths distant from the mouth opening.

The buccal tube has rather thin walls, forming about 10% of the

proboscis diameter. It is lined with a tall folded epithelium.

Buccal mass and oesophagus

The buccal mass is large in comparison with the proboscis, equiva-

lent to about two thirds of its length. It lies posterior to the proboscis,

with a wide lumen and relatively thin walls. Buccal lips are absent.

The oesophagus is greatly elongated between the buccal mass and

nerve ring, forming a long loop. Anteriorly, it is wide and flattened,

but soon becomes very narrow. There is no buccal sac.

Glands

The salivary glands are large, acinous and fused. The venom gland

changes in histology abruptly after passing anteriorly through the

nerve ring. The duct is very narrow, unciliated, and opens into the

buccal cavity at the posterior border with the oesophagus. The

muscular bulb is medium sized, of an irregular oval shape, with its

wall consisting of a thick outer layer of circular muscle fibres, a

connective tissue layer, a thinner layer of circular fibres and a thin

innermost layer of longitudinal fibres.

Odontophore and radula

The odontophore is medium-sized, with the odontophoral cartilages

Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

Fig. 18 Radulae of Epidirona and Gemmula (outgroup). a, Epidirona gabensis scale bar = 204m b, Gemmula deshayesi scale bar = 25um.

paired unfused. The radula is composed of marginal teeth. These

teeth (Fig.18a) are of the wishbone type with the distal half tapering

to a sharp point, whilst the proximal part of the tooth bifurcates into

two more or less equisized limbs. The marginal tooth are medium

long, ca.120um (0.5% of SL, 1.8% AL).

Inquisitor latifasciata (Sowerby, 1870)

(Figs 17b, 20)

See also Taylor (1994, plate le,7; figs 11-12)

Rhynchodeum and proboscis

The rhynchodeal sphincter is large and anteriorly situated. The

epithelium of the anterior two thirds of the rhynchodeal cavity

consists of very tall cells, is glandular, and folded. The epithelium of

the posterior one third of the rhynchodeum is continuous with that of

the proboscis wall, and consists of low, cubic, non-glandular cells.

The rhynchostome is narrow.

The proboscis is long in semi-relaxed animals, and lies coiled

within the rhynchodeal cavity. In retracted specimens, the proboscis

is very short, less that one third of the rhynchocoel and infolded at

the tip. The proboscis wall is not thick in retracted specimens, and

comprises about 8% of the proboscis diameter. The mouth is narrow

in semi-relaxed animals, but is capable of great stretching and is very

wide in retracted specimens (Fig. 20). The muscles of the proboscis

wall are equally developed along its length. Both anterior and

intermediate buccal tube sphincters are absent and there is no sac-

like enlargement of the buccal tube. The buccal tube is very

thin-walled in the anterior part (in the inverted position of the

proboscis tip), but rather thick posteriorly, where it is nearly equal to

the proboscis wall in width. The buccal lips are very large and form

the muscular tube, which in retracted specimens extends beyond the

mouth of the proboscis.

Buccal mass and oesophagus

The buccal mass is muscular, with thick walls and a rather narrow

inner cavity which is not curved. In the retracted state, it lies just

within the base of the proboscis, but can clearly be protracted to near

the distal tip of the proboscis. There are several retractor muscles,

attached to the buccal mass at one end and to the rhynchodeum at the

other. The oesophagus is slightly elongated between the buccal mass

and nerve ring, forming a short loop.

Glands

The salivary glands are large, paired and acinous. The histology of

the venom gland changes abruptly in histology after passing ant-

eriorly through the nerve ring. The duct of the gland is narrow and

opens just posterior to the buccal cavity. The muscular bulb wall is

formed of two equal layers of longitudinal muscle fibres, divided by

a connective tissue layer.

Odontophore and radula

The odontophore is medium sized, with paired, unfused,

odontophoral cartilages. The buccal sac is long. The radular com-

prises marginal teeth only, with each tooth of the wishbone type

(Fig. 17b) with a large robust major limb with a distal point and a

smaller, slender secondary limb

Inquisitor aemula (Angas, 1877)

(Figs 17c, 21)

Rhynchodeum and proboscis

The rhynchodeal sphincter is medium-sized and anteriorly located.

The epithelium of nearly the whole rhynchodeum is glandular and

only in a small posterior portion is it low, cubic and non-glandular,

like that of the proboscis.

The proboscis is very long, longer than the rhynchodeal cavity

and curved when retracted. The proboscis walls are thin, comprising

about 10% of the proboscis diameter. The mouth is very narrow, with

a low ‘rim’. Muscles of the proboscis wall are equally developed

along its length. There is no anterior buccal tube sphincter, but there

is a large intermediate sphincter, which lies at a distance of about

three marginal tooth lengths from the mouth opening. The sac-like

enlargement of the buccal tube has a wide lumen, with the epithe-

lium near the mouth opening being columnar and moderately tall,

soon becoming low, but forming the epithelial pad, to which a

marginal tooth was attached. The buccal tube has moderately thick

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS 73

Cerin

Fig. 19 Funa jeffreysii (Smith, 1875). A, Semidiagrammatic longitudinal section of the foregut; B, longitudinal section of the proboscis tip with a gripped

marginal tooth.

Fig. 20 Inquisitor latifasciata (Sowerby, 1870). Longitudinal section of the proboscis and buccal mass in the retracted position.

walls, which are equal in thickness to the proboscis walls. It is wide Buccal mass and oesophagus

after the intermediate sphincter, but then greatly reduced in diam- The buccal mass lies within the proboscis and occupies about half

eter, becoming wider posteriorly (on the drawing the narrowed part the proboscis length. It has rather thin walls and a broad inner cavity,

of the buccal tube is somewhat shorter, than actual). The buccal lips which shows no curvature. The oesophagus is greatly elongated

are small. between the buccal mass and nerve ring, forming a long loop.

aoe IPI Ig

Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

gre bip

Fig. 21. Inquisitor aemula (Angas, 1877). A, semidiagrammatic longitudinal section of the foregut (salivary ducts not shown); B, longitudinal section of

the proboscis tip.

Glands

The salivary glands are medium-sized, paired and acinous, with the

ducts thick and coiled. The venom gland changes abruptly in histol-

ogy after passing anteriorly through the nerve ring. The duct of the

gland is unciliated, narrow, with thick muscular walls, and opens just

posterior to the buccal cavity. The muscular bulb is formed of an

outer layer of longitudinal muscle fibres, a connective tissue layer,

an inner layer of longitudinal fibres, which is ca. 2.5 times thinner

than the outer and finally, a thin, innermost layer of circular muscle

fibres. The epithelium lining the inner cavity is rather well devel-

oped and non-glandular.

Odontophore and radula

The odontophore is medium-sized, with the odontophoral cartilages

paired and unfused. The buccal sac is medium long and very narrow.

The radula (Fig. 17c) consists of marginal teeth of the wishbone

type, with a robust, solid, pointed, major limb and a smaller thinner,

minor limb. The marginal tooth is medium long, ca.130um (0.6% of

SL, 2.3% AL).

Inquisitor aff. adenicus Sysoev,1996

(Fig. 17d)

Rhynchodeum and proboscis

The rhynchodeal sphincter is medium large and anteriorly located.

The epithelium of nearly whole the rhynchodeum is glandular and

for only about one quarter of the posterior portion is it continuous

with that of the proboscis wall.

The proboscis is long, and occupies the entire rhynchocoel in one

specimen and about half of it in the other. The proboscis walls are

thick, and compose about 25% of proboscis diameter. The mouth is

very narrow and lined with probable sensory epithelium. The mus-

cles of the proboscis wall are equally developed along its length. The

anterior buccal tube sphincter is very small and hardly visible, but

the intermediate buccal tube sphincter is large, and lies at a distance

of more than two marginal tooth lengths behind the mouth opening.

The sac-like enlargement of the buccal tube is well developed, and

lined with tall epithelium, which in one specimen forms a pad with

a marginal tooth attached to it. The buccal tube has thin walls, about

34% of proboscis diameter, lined with a tall epithelium. The buccal

lips are very small.

Buccal mass and odontophore

The buccal mass is short, about one third of proboscis length and lies

posterior to the base of the proboscis. It is uncurved, with rather thin

walls and a broad inner cavity. Several folds of the walls project into

the buccal cavity. The buccal mass lies posterior to the proboscis

base. The oesophagus is greatly elongated between the buccal mass

and nerve ring, forming a long loop.

Glands

The salivary glands are acinous, with thick, uncoiled ducts. The

venom gland changes abruptly in histology after passing through the

nerve ring. The duct of the gland is narrow and unciliated, with thick

muscular walls, and opens just posterior to the buccal cavity. The

muscular bulb is large and long, with its wall formed of two nearly

equal layers of circular muscle fibres, divided by a connective tissue

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

layer. The epithelium, lining the inner cavity is rather well-devel-

oped in one specimen, but non-glandular.

Odontophore and radula

The odontophore is medium-sized with paired, unfused odontophoral

cartilages. A buccal sac is absent. The radula (Fig. 17d) consists of

marginal teeth which are of the wishbone type with a robust major

limb and a smaller secondary limb. The marginal tooth is short,

ca.140um (0.5% of SL, 1.4% AL).

Funa jeffreysii (Smith, 1875)

(Figs 19, 23d)

See also Taylor (1994, plates If, 6a; figs 8 & 9)

Rhynchodeum and proboscis

The rhynchodeal sphincter is large and anteriorly located. The

epithelium of the anterior one third of the rhynchodeum is highly

folded and glandular, formed of large cells. In the posterior of the

rhynchodeum there is an abrupt change to a very low and incon-

spicuous, non-glandular epithelium which is continuous with that of

the proboscis wall.

The proboscis is long, as long as the rhynchodeum, with the tip

not infolded. The proboscis walls are thin, comprising about 7% of

proboscis diameter. The walls of the buccal tube are similarly thin,

composing about 5% of total diameter. The mouth is narrow. The

muscles of the proboscis walls are equally developed along its

length.

There is a medium-sized, anterior buccal tube sphincter, posi-

tioned at the base of the sac-like enlargement of the buccal tube. An

epithelial pad is present within the sac-like enlargement. There is no

intermediate buccal tube sphincter. The buccal tube is lined with a

very low epithelium. It expands greatly to form a long circular fold,

which appears like poorly-muscularized, anteriorly-directed lips,

similar to the ‘valvule’ (Sheridan ef al., 1973) The buccal lips are

large and muscular.

Buccal mass and oesophagus

The buccal mass is large and situated partially within the proboscis

base. Its walls are moderately thick. The oesophagus is greatly

elongated between the buccal mass and nerve ring, forming a long

loop. The walls of the oesophageal loop are thick and formed from

distinct longitudinal muscle fibres. The buccal sac is not defined.

Glands

The salivary glands are large, consisting of single tubes, surrounded

by acinous cells (acinous tubular type). The ducts are thick and

coiled. The venom gland changes in histology while passing

anteriorly through the nerve ring. The duct of the gland is narrow,

unciliated and opens into the posterior buccal cavity. The muscular

bulb is medium-sized, its wall formed of two layers of circular

muscle fibres (the outer layer being nearly three times thicker than

the middle, divided by a connective tissue layer, with a third inner-

most, thin layer of longitudinal muscle fibres.

Odontophore and radula

The odontophore is medium-sized, with paired, unfused cartilages.

The radula consists of marginal teeth (Fig. 23d which are paddle-

shaped with a broad distal blade and a long narrow shaft. The distal

tip is pointed, with knife-like edges on either side and a blunt barb.

An inconspicuous, thin, secondary limb lies along the margin of the

main shaft of the tooth.

Remarks

The unusual feature of this species is that the buccal mass can be

protracted way beyond the proboscis tip and out through the

rhynchostome (Taylor, 1994, fig. 8). Nevertheless, in the retracted

position the buccal mass lies at the proboscis base. It is possible that

the presence of the valvule is also connected with the possibility of

buccal mass eversion.

Funa latisinuata (Smith, 1877)

(Fig. 23c)

See also Taylor (1994, plate 1g, 6b; fig.10)

Rhynchodeum and proboscis

The rhynchodeal sphincter is large. The epithelium of the anterior

one third of the rhynchodeum is highly folded and glandular whilst

the posterior is composed of a low, inconspicuous, non-glandular

and continuous with that of the proboscis wall.

The proboscis is long, nearly as long as the rhynchodeum and

broad with the tip not infolded. The proboscis and buccal tube walls

are thin. The mouth is relatively wide. The muscles of the proboscis

wall are equally developed along its length. There is a small anterior

buccal tube sphincter, but no sac-like enlargement, no epithelial pad

and no intermediate sphincter.

Buccal mass and oesophagus

The buccal mass is large and situated partially within the proboscis

base and uncurved. Its walls are moderately thick and muscular. The

oesophagus is greatly elongated between the buccal mass and nerve

ring forming a long loop. The walls of the oesophageal loop are thick

and muscular. There is no buccal sac.

Glands

The salivary glands are large and acinous, but with a modified

histology. The ducts are thick and coiled. The venom gland changes

in histology while passing anteriorly through the nerve ring. The

anterior part has thick muscular walls and opens into the posterior

part of the buccal cavity. The muscular bulb is medium-sized, with

the wall formed from two equisize layers of circular muscles,

divided by a thick connective tissue layer, with a very thin innermost

layer of longitudinal muscle fibres.

Odontophore and radula

The odontophore is medium-sized, with paired, unfused cartilages.

The radula consists of marginal teeth only (Fig. 23c) which are

paddle-shaped, with a flattened, angular, barb-less, distal termina-

tion, and a long thin shaft. A thin accessory limb lies along the shaft.

Ptychobela suturalis (Gray, 1838)

(Figs 22, 23a, 30c)

Rhynchodeum and proboscis

The rhynchodeal sphincter is medium-sized and situated rather to

the posterior of the rhynchostome. The epithelium of the anterior

one third of the rhynchodeal cavity is folded and formed of tall,

glandular cells. The low, non-glandular epithelium of the posterior

two thirds of the rhynchodeum is continuous with that of the

proboscis wall. This indicates that a large part of the rhynchodeal

wall takes part in proboscis protraction.

In retracted animals, the proboscis is short with the tip infolded

(Taylor 1994, fig. 18). However, in relaxed animals, the proboscis is

very long, longer that the rhynchocoel and with the tip not inverted.

Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

Fig. 22. Ptychobela suturalis (Gray, 1838). A, semidiagrammatic longitudinal section of the foregut (only one salivary duct shown) with the proboscis in

an everted position and buccal lips inverted inside the buccal cavity; B, longitudinal section of the proboscis tip.

The proboscis walls are medium-thick, and comprise about 15%

of the proboscis diameter. The mouth is narrow in relaxed animals,

but capable of great expansion. The muscles of the proboscis walls

are equally developed along its length.

The buccal tube has a very small anterior sphincter and there is no

sac-like enlargement at the distal end There is large intermediate

sphincter, which lies about two radular tooth lengths behind the

proboscis tip. The buccal tube has rather thick walls, only slightly

thinner than those of the proboscis. In the anterior half of the

proboscis, the buccal tube is narrow, but greatly expanded posteriorly.

There are large, extensible buccal lips which can invert into the

buccal cavity. In retracted specimens, they form a muscular tube

with a flaring aperture, which extends beyond the mouth of the

proboscis.

Buccal mass and oesophagus

The buccal mass lies within the base of the proboscis, and is rather

long, comprising about a quarter of the proboscis length, with

relatively thin walls and a very broad inner cavity, which is not

curved. The oesophagus is elongated between the buccal mass and

nerve ring, forming the short loop.

Glands

The salivary glands are large, paired and acinous. The venom gland

changes in histology after passing anteriorly through the nerve ring.

The duct of the gland is narrow, ciliated, and opens into posterior

part of the buccal cavity. The muscular bulb is long, with the wall

formed of two equally thick layers of longitudinal muscle fibres,

divided by a connective tissue layer and very thin innermost layer of

circular fibres.

Odontophore and radula

The odontophore is small with the odontophoral cartilages, paired,

and unfused. The buccal sac is very short. The radular teeth (Fig.

23a) are hollow and awl-shaped with a sharp point and broadening

towards the base. They are composed of two separate pieces which

are fused along one edge and loosely twisted together.

Cheungbeia mindanensis (Smith, 1877)

(Figs 24, 26d, 30d)

See also Taylor (1994, plates 1h, 6d; figs 13-15)

Rhynchodeum and proboscis

The rhynchodeal sphincter is large and anteriorly located and the

rhynchostome is relatively wide. The epithelium of the anterior half

of the dorsal wall of the rhynchodeal cavity is glandular (dorsal wall

is significantly longer than the ventral). In the posterior part of the

rhynchodeum the epithelium is non-glandular and continuous with

that of the proboscis wall. The proboscis is extremely long and lies

coiled within the rhynchocoel. The proboscis walls are highly folded

and comprise about 15% of the proboscis diameter. The muscles of

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS 7

Fig. 23. Radulae of Crassispirinae. a, Ptychobela suturalis scale bar = 20pm b, Vexitomina garrardi scale bar = 20um c, Funa latisinuata scale bar = 50um

d, Funa jeffreysii scale bar = 60um.

vga

Fig. 24 Cheungbeia mindanensis (Smith, 1877) A, longitudinal section of the proboscis tip, the tooth at the same scale shown below; B, longitudinal

section of the proboscis base and buccal mass (salivary ducts not shown).

78 Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

Fig. 25 Cheungbeia robusta (Hinds, 1843). A, longitudinal section of the proboscis tip, with a tooth shown at the same scale below; B, longitudinal

section of the proboscis base and buccal mass.

Fig. 26 Radulae of Cheungbeia. a—c, Cheungbeia robusta, a, portion of radula with two rows of teeth, scale bar = 30um b, detail of tooth tip scale bar =

10um c, single tooth, note secondary limb (arrowed) along edge of major limb scale bar = 20um d, Cheungbeia mindanensis single tooth with secondary

limb arrowed. scale bar = 25um.

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

the proboscis wall are equally developed along its length. The mouth

is narrow. The proboscis is covered with ciliated epithelium with

long cilia, which is replaced by non-ciliated epithelium with a rather

thick cuticle at its base.

A small, anterior, buccal tube sphincter lies in front of the sac-like

enlargement of the buccal tube, close to mouth opening. The epithe-

lium, lining the enlargement is tall, forming pads on both walls (Fig.

30d). A second larger sphincter lies at the base of the enlargement at

the distance slightly longer than one tooth length and therefore

should be considered as anterior. The walls of the buccal tube are

thin in the anterior part, but become rather thick (about 10% of

proboscis diameter) after the second sphincter.

Buccal mass and oesophagus

The buccal mass lies at the proboscis base and is short in comparison

to the proboscis. It is thick-walled, with a broad lumen and curved.

There are very large, extensible buccal lips. The oesophagus is

greatly elongated between buccal mass and nerve ring and forms a

long loop. There is no buccal sac. The opening of the radular

diverticulum is narrow and relatively long. The salivary ducts open

into the buccal cavity on both sides of the opening of the radular

diverticulum.

Glands

The salivary glands are medium-sized, paired and acinous. The

venom gland changes markedly in histology after passing through

the nerve ring. The duct is very narrow, unciliated and opens at the

border between the buccal mass and oesophagus. The muscular bulb

is medium-sized, with the wall formed from two layers of longitudi-

nal muscle fibres (the outer about twice as thick as the inner),

divided by a connective tissue layer, with a thin, innermost layer of

circular muscle.

Odontophore and radula

The odontophore is rather large, with paired, unfused cartilages. The

radula consist of marginal teeth (Fig. 26d), which are long and

harpoon-like in form, but concavo-convex (gutter-shaped) in profile.

The distal end of the tooth is sharply pointed with a pronounced

barb, whilst the shaft is more or less straight-sided, with just a slight

enlargement at the base. A very thin secondary limb lies along the

edge of the shaft (Fig. 26d). The marginal tooth is very long, ca.

320um (1.3% of SL, 4.0% AL).

Cheungbeia robusta (Hinds, 1843)

(Figs 25, 26a—c)

See also Taylor (1994, plates li, 6c; figs 16-17)

Rhynchodeum and proboscis

The rhynchodeal sphincter is large and located slightly to the

posterior. The epithelium of nearly the whole wall of the rhynchodeal

cavity is glandular, whilst only a small posterior portion is non-

glandular and continuous with that of the proboscis wall. The

rhynchostome is rather wide.

The proboscis is very long, longer than the rhynchocoel and

coiled within it. It is relatively wide at the base and narrows towards

the tip. The walls of the proboscis are thin and highly folded,

forming about 10% of the proboscis diameter. Muscles of the

proboscis wall are equally developed along its length. The mouth is

narrow. The anterior, buccal tube sphincter is very small and lies

close to the mouth opening, in front of the short, only slightly

differentiated, sac-like enlargement of the buccal tube The epithe-

lium lining this enlargement is tall, forming pads on both walls. A

second, larger sphincter lies at the base of the enlargement at slightly

more than one radular tooth length from the mouth and therefore

should be considered as an anterior sphincter. The buccal tube is

narrow anteriorly, but expands greatly posterior to the sac-like

enlargement and occupies nearly the whole proboscis lumen. Never-

theless, its walls are thin and form only about 5% of the proboscis

diameter at its base.

Buccal mass and oesophagus

The buccal mass is long, thick-walled with broad lumen, curved, and

lies at the proboscis base. The boundary between the posterior

buccal tube and the buccal mass is not well defined. The buccal lips

are very short. The oesophagus is greatly elongated between buccal

mass and nerve ring and forms a long loop, which is narrow when

leaving the buccal cavity and expands posteriorly. There is no buccal

sac. The opening of the radular diverticulum is rather narrow and

moderately long. The thick and long salivary ducts open in the

buccal cavity on both sides of the opening of the radular diverticu-

lum.

Glands

The salivary glands are medium-sized, paired, and of the modified

acinous type. The venom gland changes abruptly in histology after

passing through the nerve ring. The duct is very narrow, unciliated

and opens at the border between the buccal mass and oesophagus.

The muscular bulb is large, with the wall mainly formed of two

layers of circular muscle fibres (the outer being about twice the

thickness of the inner), divided by a connective tissue layer, with a

thin, innermost layer of circular muscle.

Odontophore and radula

The odontophore is small, with paired, unfused cartilages. The

radula consist of marginal teeth only (Figs 26a—c) which are har-

poon-like and very similar to those of Cheungbeia mindanensis

(Fig. 26d). These have long shafts which are concavo-convex (gutter

shaped) in profile. The distal end of the tooth is sharply pointed with

a pronounced barb, whilst the shaft is more or less straight-sided,

with just a slight enlargement at the base. A very thin secondary limb

lies along the edge of the shaft (Fig. 26c).

Antiguraleus morganus (Barnard, 1958)

(Figs 27, 28a)

Rhynchodeum and proboscis

The rhynchodeal sphincter is medium-sized and anteriorly located.

The epithelium of the anterior half of the rhynchodeal cavity is tall,

glandular, folded and formed of large cells, while that of the poste-

rior half of the rhynchodeum is non-glandular and continuous with

that of the proboscis wall. This posterior part of the rhynchodeum is

attached by numerous muscle fibres to the body wall and probably

cannot be everted. The rhynchostome is narrow.

The proboscis is very long, more than twice as long as the

rhynchodeal cavity, rather thin and coiled. The proboscis walls

comprise about 15% of the proboscis diameter. The mouth is very

narrow. The muscles of the proboscis wall are equally developed

along its length. The anterior buccal tube sphincter is small, and

lies in front of a very broad sac-like enlargement of the buccal

tube. The epithelium, lining the enlargement is similar to that of

the rest of the buccal tube. Additionally, there is large intermediate

sphincter, situated approximately half-way along the proboscis.

The buccal tube has rather thin walls, forming about 8% of probos-

cis diameter.

con

btsa = ebt ngre btsi

Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

gre

rhs

LUT

ma ;

a, Wad

Fig. 28 Radulae of Crassispirinae. a, Antiguraleus morganus b, Paraguraleus costatus. Scale bars = 20um.

Buccal mass and oesophagus

The long, buccal mass lies posterior to the proboscis, and is equiva-

lent to nearly half of the proboscis length, with thick walls and a

rather broad inner cavity, which is not curved. There are large

extensible buccal lips. The oesophagus is not elongated between

buccal mass and nerve ring, which is situated closely posterior to the

buccal mass. A buccal sac is absent.

Glands

The salivary glands are large, paired, and consist of single, coiled

tubes but with the acinous morphology. The venom gland does not

change histology after passing anteriorly through the nerve ring. It

opens at the border between the buccal mass and oesophagus just in

front of the nerve ring. The muscular bulb is small, the wall formed

of two equal layers of longitudinal fibres, divided by a connective

tissue layer, with an innermost very thin layer of circular muscle.

Odontophore and radula

The odontophore is medium-sized, with a pair of unfused cartilages

formed of one layer of cells. The radula consists of marginal teeth of

the wishbone type (Fig. 28a). The major limb of the tooth is elongate

with a pointed tip and a constricted waist in the middle of the tooth.

Below this constriction, the lower marginal edge of the tooth is

extended as a ‘soft’ buttress to attach to the radular membrane. The

minor element is slender and shorter, attached to the distal blade of

the major element and broadens slightly at the base, where it

attaches to the membrane. The marginal teeth are long, ca.120um

(1.2% of SL, 3.2% AL).

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

Guraleus costatus (Hedley, 1922)

(Fig. 28b)

Unfortunately, the specimen was sectioned nearly transversely and

an illustration of it comparable with the other species was not

possible.

Rhynchodeum and proboscis

The rhynchodeal sphincter is small and anteriorly located. The

epithelium of the anterior part of the rhynchodeal cavity is glandular,

whilst that of the posterior rhynchodeum is nonglandular and con-

tinuous with that of the proboscis wall. The non-glandular posterior

part of the rhynchodeum is very thin and attached by numerous

muscle fibres to the body wall and probably cannot be everted. The

rhynchostome is narrow.

The proboscis is very long and coiled, more than twice as long as

the rhynchodeal cavity. It is very thick at the base but narrows

towards the tip (diameter of 0.44 mm at the base, but only 0.08 at the

tip), The proboscis walls are thin, comprising less than 7% of the

proboscis diameter at its base. The mouth is very narrow. A very

small, anterior buccal tube sphincter, lies in front of a small, sac-like

enlargement of the buccal tube. The epithelium, lining the enlarge-

ment is similar to that of the rest of the buccal tube. A single tooth

was seen in the enlargement. There is large intermediate sphincter,

situated approximately midway down the proboscis. The buccal

tube is very narrow anteriorly, but broad in the posterior part of the

proboscis. Its walls are very thin and highly folded.

Buccal mass and oesophagus

The buccal mass lies within the base of the proboscis, with thick

walls and rather broad inner cavity, and showing no curvature. There

are large, extensible, buccal lips, which can be inverted inside the

cavity. The oesophagus is slightly elongated between buccal mass

and nerve ring and forms a short loop. There is no buccal sac. The

salivary ducts open into the buccal cavity on both sides of the

opening of the radular diverticulum.

Glands

The salivary glands are small, paired and acinous. The venom gland

changes abruptly in histology after passing through the nerve ring.

The duct is very narrow, unciliated and opens at the border between

the buccal mass and oesophagus. The muscular bulb is large, the wall

is formed of two equal layers of longitudinal fibres, divided by a

connective tissue layer, with a thin, innermost layer of circular

muscle.

Odontophore and radula

The odontophore is rather large with a pair of unfused cartilages.

The radula consists of marginal teeth of the wishbone type (Fig.

28b). Each tooth has a robust and pointed major limb with a straight,

blade-like leading edge. The secondary limb is long and slender and

attached to the major limb near the distal tip. The marginal teeth are

very long, ca.105um (1.4% of SL, 4.2% AL).

Burchia spectabilis Sysoev & Taylor, 1997

(Fig. 29)

See Sysoev & Taylor (1997, fig. 3)

Rhynchodeum and proboscis

The rhynchodeal sphincter is very large, and located slightly to the

posterior. The anterior one third of the rhynchodeal cavity possesses

a very tall, glandular, folded epithelium formed of large cells. The

epithelium of the posterior two thirds of the rhynchodeum is con-

tinuous with that of the proboscis wall and is low, cubic and

non-glandular. This indicates, that the greater part of the rhynchodeal

lining is involved in proboscis protraction. The rhynchostome is

narrow.

asgm

Fig. 29 Burchia spectabilis Sysoev & Taylor, 1997. Semidiagrammatic longitudinal section of the foregut (salivary ducts not shown).

82 Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

Fig. 30 a, Nquma scalpta, longitudinal section of foregut showing proboscis and glandular anterior part rhynchodeal wall. Scale bar = 250um b,

Vexitomina garrardi showing proboscis and muscular wall of the posterior rhynchodeum continuous with proboscis wall and also the radular sac located

some distance to the posterior of the proboscis base. Scale bar = 250um c, Ptychobela suturalis showing large buccal lips, inverted into the buccal cavity.

Scale bar = 250um d, Cheungbeia mindanensis, sac-ike enlargement of the buccal tube near the proboscis tip showing the pad of tall epithelial cells.

Scale bar = 60um e, Crassispira harfordiana flucki, long proboscis in the rhynchocoel showing the radular sac located in the anterior part of the

proboscis and the complex folding of the proboscis tip (see Fig. 6). Scale bar = 250um f, longitudinal section of the proboscis tip of the outgroup

Gemmula deshayesi (Turrinae) showing the wishbone tooth held in the sac-like enlargement of the buccal tube and the intermediate sphincter located

several tooth lengths from the proboscis tip. Scale bar = 250m.

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

The proboscis is long and thick, and occupies about two thirds of

the rhynchodeal cavity. The proboscis walls are thick and comprise

about 20% of the total diameter. The mouth is narrow in the

preserved condition, but appears capable of great enlargement. The

ventral side of the anterior part of the proboscis wall is invaginated.

The muscles of the proboscis wall are equally developed along its

length. Both anterior and intermediate buccal tube sphincters are

absent, as is also the distal sac-like enlargement of the buccal tube.

Buccal mass and oesophagus

The buccal mass is situated to the posterior of the base of the

proboscis. It is long, equivalent to about half of the proboscis length,

with thick walls and a rather narrow inner cavity, which is not

curved. Extensible buccal lips are absent. The oesophagus is elon-

gated between the buccal mass and nerve ring, forming a short loop.

Glands

The salivary glands large and acinous. The venom gland shows a

change in histology after passing anteriorly through the nerve ring.

The duct of the gland is narrow, ciliated and opens into the posterior

part of the buccal cavity. The muscular bulb is long with the wall

formed of two layers of equal thickness composed of longitudinal

fibres, divided by a connective tissue layer.

Odontophore and radula

The odontophore is medium-sized, with paired, unfused,

odontophoral cartilages formed of one layer of cells. The radula

comprises marginal teeth, which are of the robust, wishbone form,

with a long solid, distally pointed major limb and a thinner second-

ary limb, which attaches near the tip of the major limb. The marginal

teeth are short, ca. 165um (0.5% of SL, 1.4% AL).

Vexitomina garrardi (Laseron, 1954)

(Figs 23b, 30b)

Rhynchodeum and proboscis

The rhynchodeal sphincter is large, long and situated slightly to-

wards the posterior. The epithelium of the anterior two thirds of the

rhynchodeum is glandular, forming tall folds, whilst the epithelium

of the posterior one third of the rhynchodeum is non-glandular and

continuous with that of the proboscis wall. The rhynchostome is

wide. The proboscis is short, cone-shaped, and occupies about half

the rhynchocoel. The proboscis walls form about 20% of the probos-

cis diameter. The muscles of the proboscis walls are equally developed

along its length. The mouth is very narrow.

The anterior buccal tube sphincter is small, and lies close to the

mouth, opening in front of the long sac-like enlargement of the

buccal tube. A tall epithelium lines the enlargement. A second, larger

sphincter lies at the base of the enlargement, at a distance slightly

longer than one tooth length and therefore should be considered as

anterior. A tooth was seen in the buccal tube posterior to the latter

sphincter. The buccal tube walls are rather thin (about 8% of

proboscis diameter) and highly folded. In the posteriormost part the

tube forms a very long, but narrow, circular fold.

Buccal mass and oesophagus

The buccal mass is large in comparison with the proboscis, equiva-

lent to about two thirds of its length, with thick, folded walls and a

narrow lumen. It lies posterior to the proboscis base. The buccal lips

are small. The oesophagus is elongated between the buccal mass and

nerve ring and forms a short loop. After leaving the buccal cavity, the

oesophagus is very narrow, but then expands greatly after passage

through the nerve ring. The buccal sac is very short.

Glands

The salivary glands are medium-sized, paired, and acinous. The

venom gland changes sharply in histology after passing anteriorly

through the nerve ring. The duct is very narrow, unciliated and opens

at the border between the buccal mass and oesophagus.

The muscular bulb is medium-sized, with the wall formed of two

layers of longitudinal muscle fibres (the outer being twice as thick as

the inner), divided by a connective tissue layer and innermost thin

layer of circular muscle.

Odontophore and radula

The odontophore is medium-sized, with paired, unfused cartilages.

The radula consists of marginal teeth (Fig. 23b) which are long, with

a blade-like distal portion, which is sharply pointed with a small

barb. The shaft is long and tapers gradually towards the base. The

teeth are slightly concavo-convex in profile and a very thin splint-

like secondary limb lies along the edge of the shaft.

Turridrupa bijubata (Reeve, 1843)

Rhynchodeum and proboscis

The rhynchodeal sphincter is medium-sized and posteriorly located.

The anterior half of the rhynchodeum has a highly folded, tall,

glandular epithelium whilst the posterior half has a low cubic

epithelium similar to that of the proboscis wall. The proboscis is

short and occupies about half of the rhynchodeum, the tip is thin,

while the base is thicker and muscular. There is a very small anterior

buccal tube sphincter, lying at the distal end of a small sac-like

enlargement containing a single radular tooth. A larger intermediate

sphincter lies at a distance of about 10 tooth lengths from the

proboscis tip.

Buccal mass and oesophagus

The buccal mass is long and curved and lies to the posterior of the

proboscis. The walls of the buccal mass are long and muscular

anterior to the entrance of the radular sac and the buccal lips small

and not invertible. The oesophagus is not elongated between the

buccal sac and nerve ring.

Glands

The salivary glands are large, paired and acinous, with large ciliated

ducts. The venom gland changes abruptly in histology after passing

anteriorly through the nerve ring and the duct opens into the oesopha-

gus just posterior to the buccal mass. The muscular bulb is large and

comprises two layers of circular muscle separated by a connective

tissue layer. The outer layer is about twice as thick as the inner. There

is no thin, innermost, muscular layer.

Odontophore and radula

The odontophore is large with two large unfused cartilages. The radula

consist of both central and marginal teeth (Kilburn, 1988, fig. 40).The

central tooth comprises a square plate with a prominent spine-like

cusp. The marginal teeth are of the robust wishbone type with the

bifurcating proximal end, similar to Epidirona gabensis (Fig. 18a).

Outgroup

This species from the family Turrinae, was chosen as the outgroup.

The anatomy and radula is described and illustrated in Taylor (1994)

and further details are given below.

Gemmula deshayesii (Doumet, 1839)

(Figs 18b, 30f )

Rhynchodeum and proboscis

There is a large rhynchodeal sphincter situated slightly to the

posterior. The rhynchodeum has a folded epithelium of tall glandular

cells for its entire length.

The proboscis is long, nearly as long as the rhynchodeum. The

proboscis walls are medium thick about 15% of total diameter,

whilst the walls of the buccal tube are thin forming about 5% of the

diameter. The muscles of the proboscis wall are less developed near

the tip. There is a small anterior buccal tube sphincter, with a short

sac-like enlargement and epithelial pad. A single wishbone radular

tooth was held in the enlargement. An large intermediate buccal tube

sphincter lies at about five tooth lengths from the mouth.

Buccal mass and oesophagus

The buccal mass is short, muscular and uncurved and lies within the

base of the proboscis. The buccal lips are medium long and invert-

ible. The oesophagus is not elongated between the nerve ring and

buccal mass.

Glands

Salivary glands are large and acinous with paired ducts. There is no

change in the histology of the venom gland to the anterior of the

nerve ring. The gland opens into the rear of the buccal cavity by a

short ciliated duct. The muscular bulb comprises two subequal

layers of circular muscle divided by a connective tissue layer.

Odontophore and radula

There a two medium-sized odontophoral cartilages. The buccal sac

is medium long. The radula consist of both central and marginal

teeth (Fig. 18b) The central tooth is wide and low with a central

spine-like cusp. The marginal teeth are wishbone in form, but of the

‘clothes-peg’ type with pointed tips and bifurcated proximal ends.

CHARACTER ANALYSIS AND RELATION-

SHIPS WITHIN THE CRASSISPIRINAE

Characters and states

On the basis of the analysis of the thin sections described above and

previous work (Taylor et al., 1993) we selected 20 characters with

potential for determining relationships between the species studied.

These characters and their states are listed in Table 2. Many of these

are self explanatory or reference is given to figures which illustrate

the various states. However, further explanation of some characters

is given below.

Character 2. In some species, such as Inquisitor latifasciata and

Ptychobela suturalis, the proboscis tip may be inverted deeply

into the buccal tube (Fig. 20).

Character 5. In some species there is an epithelial pad of large cubic

cells in the anterior part of the buccal tube (C. (Gibbaspira)

dysoni, Fig. 5).

Character 7. In many species there is a sac-like enlargement of the

anterior part of the buccal tube. This is often lined with an

epithelium of tall cells. The function of the sac is to hold single

detached radular teeth at the proboscis tip (Fig. 30d).

Character 9. Some taxa possess large buccal lips which are often

capable of inversion into the buccal cavity; in others this was not

observed.

Character 11. In many species there is a sharp bend within the

buccal mass (e.g. Fig. 9), while other species show no curvature.

Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

Table 2 List of characters and states used in the cladistic analysis

1. Epithelium of posterior rhynchodeal wall: 0 — glandular;

1 —continuous with proboscis wall for less than 12 of the

rhynchodeum length; 2 — continuous with proboscis wall for more

than 2 of the rhynchodeum length.

2. Proboscis tip: 0 — not inverted inside; 1 — inverted inside.

Proboscis tip epithelium: 0 — not invaginated; 1 — invaginated.

4. Proboscis length: 0 — very long (longer than rhynchodeum);

1 — long (50-100% of rhynchodeum length); 2 — short (less than 50%

of rhynchodeum length).

Epithelial pad at tip of buccal tube: 0 — present; 1 — absent.

6. Anterior buccal tube sphincters (the distance from the sphincter to the

mouth opening is less than the radular tooth length): 0 — one; | — two;

2 — absent.

7. Sac-like enlargement of the buccal tube with tall epithelium:

0 — present; 1 — absent.

8. Intermediate sphincter of the buccal tube (the distance from the

sphincter to the mouth opening is more than the tooth length):

0 — absent; | — present.

9. Buccal lips: 0 — large invertible; 1 — large uninvertible; 2 — small;

3 — absent.

10. Position of the buccal mass: 0 — posterior to the proboscis base;

1 — at the proboscis base; 1 — within the proboscis (up to near tip).

11. Buccal mass shape: 0 — not curved; 1 — curved.

12. Elongation of the oesophagus between buccal mass and nerve ring:

0 — absent; | — present.

13. Salivary glands: 0 — acinous; 1 — modified acinous; 2 — acinous

tubular; 3 — simple tubular; 4 — anastomosing tubular.

14. Histology of the venom gland anterior to the nerve ring:

0 — unchanged; | — changed.

15. Position of opening of venom gland into oesophagus: 0 — into rear

part of buccal mass; | — into oesophagus behind buccal mass.

16. Number of muscular layers of the muscular bulb: 0-2 layers; 1-3

layers.

17. Orientation of fibres in outer two layers of muscular bulb:

0 — similar orientation; 1 — opposite orientation.

18. Raddula: central ‘tooth’: 0 — absent; 1 — central ridges present;

2 — spinose tooth.

19. Radula: curved lateral ‘teeth’: 0 — absent; 1 — present.

20. Radula: marginal teeth: 0 — Epidirona type; 1 — Inquisitor type;

2 — Funa type; 3 — Haedropleura type; 4 — Hindsiclava type;

5 — Cheungbeia type; 6 — Ptychobela type.

ee)

Character 13. During this study we found several distinct morphologies

of the salivary glands. Many species possessed the normal acinous

type found in most neogastropods. In some species, Cheungbeia

robusta and Burchia spectabilis. the appearance is of modified

acinous (State 1). Funa jeffreysii and Antiguraleus morgani have

glands consisting of single tubes surrounded by acinous cells; this

we refer to as acinous tubular (state 2). Simple tubular glands are

found in Haedropleura septangularis and Naudedrillia, these are

highly coiled in C. latizonata (State 3). Finally, in a number of taxa

the salivary glands appear to be made up of a mass of anastomosing

tubes (State 4).

Character 14. In most Crassispirinae, the venom gland becomes

ciliated and duct-like anterior to the nerve ring.

Character 17. The muscle fibres in the outer two layers of the

muscular bulb, may either have the same or differing orientations.

Character 18. Although there is no central tooth to the radula in any of

Crassispirinae studied, we did observe in two species some low

transverse ridges which cross the ribbon (Fig. 14a). We refer to

these as transverse ridges. In the outgroup, Gemmula deshayesii, a

robust central tooth with a spinose central cusp is present (Fig. 18b).

Character 19. In most crassispirines there are no lateral teeth, but

amongst the species we examined, two have paired, low, curved

structures which are symmetrical on either side of the mid-line

(Figs 11c—d). Maes (1983) referred to these structures as ‘soft

laterals’. It is uncertain whether these structures are homologous

with the lateral teeth found in the Drilliidae or other neogastropods.

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

Inquisitor Hindsiclava Haedropleura

Funa Vexitomina Cheungbeia Ptychobela

Fig. 31 Summary of major radular types found amongst the

Crassispirinae. Not to scale.

They may, as in the Columbellidae (Guralnick & de Maintenon,

1997), be produced by different secretory cells from ‘true’ teeth.

Character 20: several types of marginal teeth are found within the

taxa examined. We have divided these into 6 major types, which

are summarized in Fig. 31. The Epidirona type (State 0) (Fig. 18a)

is a wishbone tooth of the clothes-peg type with the tooth bifurcat-

ing into two more or less equal limbs. The /nquisitor type (State

1) is the most widely distributed and consists of a robust, pointed,

major limb with a smaller, slender, secondary limb attached to it.

The Funa /Vexitomina type (State 2) consists of a long tooth with

a broad, pointed blade and a narrow shaft, with a very thin

accessory limb attached to the edge of the shaft. In the

Haedropleura type (State 3), the major limb has a trowel-like

distal end and a narrow shaft, with a slender accessory limb. The

Hindisclava type (State 4) has a flat, pointed, major limb and a

long, slender, detached ‘handle-like’ accessory limb. The

Cheungbeia type (State 5) is harpoon-like with a pointed, barbed

tip and a straight concave shaft, with a thin splint-like secondary

limb attached to the edge of the shaft. Finally, the Ptychobela type

(State 6) is awl-shaped and hollow, composed of two pieces which

are fused along one edge and twisted together.

Outgroup

As the outgroup for the Crassispirinae, we chose Gemmula deshayesii

from Hong Kong, which has the relatively underived foregut anatomy

typical of members of the subfamily Turrinae. Serial sections of the

foregut of this species were already available, as well as a published

Table 3 Distribution of character states for the crassispirine gastropods

examined. Details of characters listed in Table 2.

Character

Crassispira incrassata

Crassispira maura

Crassispira dysoni

Crass. harford. flucki

Crassispira latizonata

Crassispira pluto

Crassispira kluthi

Crassispira tepocana

Crassispira xanti

Crassispira turricula

Crassispira apicata

Hindsiclava andromeda

Hindsiclava militaris

Miraclathurella bicanal.

Haedropleura septangularis

Naudedrillia prateriss.

Nquma scalpta

Epidirona gabensis

Funa latisinuata

Funa jeffreysii

Inquisitor latifasciata

Inquisitor aff. adenicus

Inquisitor aemula

Ptychobela suturalis

Cheungbeia mindanensis

Cheungbeia robusta

Antiguraleus morganus

Guraleus costatus

ide Meth Thatigdh i fai 72

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Burchia spectabilis

Vexitomina garrardi

Turridrupa bijubata

Gemmula deshayesi

description of the anatomy (Taylor, 1994). The characters of the

outgroup are recorded with the Crassispirine species in Table 3.

Character distribution

The distribution of the character states amongst the species studied

along with those of the outgroup are shown in the data matrix (Table

3). For Crassispira latizonata and Hindsiclava andromeda we had

insufficient material to investigate the radula and there are no

published descriptions. In the case of Crassispira pluto, the muscu-

lar bulb was lost from the sectioned material.

Phylogenetic analysis

This was performed using PAUP version 3.1 and McClade 3.04 for

subsequent analysis. Using the data matrix of Table 3 and the

heuristic sort option we obtained 24 equally parsimonious trees of

104 steps, with a Consistency Index of 0.33 and Retention Index of

0.54. A consensus tree generated from the 24 trees is shown in Fig.

32a and an example shown in Fig. 32b. Internal nodes of the tree are

supported by relatively few characters and there is a high level of

homoplasy. Epidirona gabensis and Turridrupa bijubata are undif-

ferentiated from the outgroup Gemmula in all trees. Whilst,

Crassispira dysoni and Inquisitor aff. adenicus which both possess

Inquisitor type teeth, are undifferentiated in the consensus tree but

form a weakly supported branch in most trees. Species classified

into different subgenera of Crassispira appear in widely separated

parts of the tree (Fig. 32b), apart from Crassispira (Crassispira) s. s.

and Crassispira (Striospira) which form a monophyletic clade.

Because we sampled only 24 out of the 46 genera and subgenera

and because of the large amount of homoplasy, it would be prema-

Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

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FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

ture to use the tree as a basis for classification of the Crassispirinae.

However, foregut characters have potential for unravelling relation-

ships amongst the gastropods with rather similar shells.

DISCUSSION

Summary of anatomical variation in

Crassispirinae

Considerable variation was found in the configuration of the foregut

amongst the species we studied. This variation is reflected in the fact

that we have recognised 13 main types of crassispirine foregut (Figs

33-34), which differ in the presence, position and morphology of the

main structures, such as buccal mass, salivary glands, buccal lips and

sphincters of the buccal tube.

Several features are characteristic for the vast majority of the

crassispirines and these include a glandular lining to the anterior part

of the rhynchodeum and the ability of the posterior part of the

rhynchodeum to evert during proboscis protraction (except

Crassispira harfordiana flucki— Fig. 33 C).This is recognised by the

continuity of the posterior rhynchodeal and proboscis walls. In

nearly all Crassispirinae, the oesophagus is elongated between the

Fig. 33. Diagram summarizing some of the major types of foregut morphology found among the Crassispirinae. Not to scale. A, Crassispira (Crassispira)

spp. — with acinous salivary glands, Funa spp. — with single tube acinous gland. B, Crassispira (Gibbaspira) dysoni. C, Crassispira (Glossispira)

harfordiana flucki. D, Crassispira (Crassiclava) spp. E, Burchia new species. F, Crassispira (Striospira) tepocana.

Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

Fig. 34 Diagram summarizing some of the major types of foregut morphology found among the Crassispirinae. Not to scale.

A, Antiguraleus morganus. B, Cheungbeia spp. C, Nquma scalpta (anastomosing tubular salivary glands) and Naudedrillia praetermissa (simple tubular

salivary glands). D, Inquisitor latifasciata and Ptychobela suturalis with a retracted proboscis. E, Ptychobela suturalis with protracted proboscis. F,

Haedropleura septangularis. G, Hindsiclava spp.

buccal mass and the nerve ring, usually forming a more or less long

loop. This character is also associated with the ability to evert the

posterior rhynchodeum. Exceptions are Antiguraleus morganus (Fig.

34 A) and Haedropleura septangularis (Fig. 34 F).

The length of the proboscis in its retracted position is very

variable, this is because the posterior part of the rhynchodeum can

evert, so that on retraction, the proboscis wall becomes the wall of

the rhynchodeum. The most extreme expression of this is found in

Hindsiclava spp. (Fig. 34 G), where the proboscis, in its retracted

position, is extremely short, occupying less that 1/5 of the rhynchocoel

length. The anatomy of the posterior part of rhynchodeum suggests

that when protracted the proboscis would be similar in length to that

of the other species.

Elongation of the oesophagus is coupled with the anterior elonga-

tion of the venom gland. This results in the formation of a usually

long, muscular, non-glandular duct, which in nearly all the

Crassispirinae studied, opens into the posterior part of the buccal

mass at the border with the oesophagus. Exceptions are Antiguraleus

morganus (Fig. 34 A) and Haedropleura septangularis (Fig. 34 F),

which are the only species lacking the oesophageal loop. In Turriinae

(Turridrupa) there is modification of the venom gland in front of the

nerve ring, but the oesophageal loop is absent.

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

In many species, there are large buccal lips, which protrude into

the buccal tube (eg. Fig. 33 A — Funa spp. 33 D — Crassispira

(Crassiclava) spp.). In several species, for example, Ptychobela

suturalis (Fig. 34 E) the buccal lips are able to invert inside the

buccal cavity. The functional significance of this feature is unknown,

but could perhaps be associated with the passage of food items into

the buccal cavity.

In four species, the tip of the proboscis is able to invert inside itself

(Figs. 33 E, 34 D). In Burchia spectabilis (Fig. 33 E), the buccal

mass is situated at the base of the proboscis and is not protruded

through the mouth, when the proboscis tip is inverted. By contrast,

the long buccal lips of Inquisitor latifasciata and Ptychobela suturalis

(Fig. 34 D) are exposed through the mouth when the proboscis tip is

inverted. In the latter species, the foregut has a totally different

appearance when the proboscis is partially protracted (Fig. 34 E).

For one species (Funa jeffreysii — Fig.19), it has been demonstrated

that the whole buccal mass can be protruded through the mouth

opening (Taylor, 1994, fig. 8). It is highly possible, that in other

crassispirines the buccal mass may also be protruded through the

mouth and used in prey capture.

Many Conoidea possesses one or more sphincters in the anterior

part of the buccal tube, these grip the detached, marginal, radular

teeth at the proboscis tip (Kantor & Taylor, 1991; Taylor et al.,

1993). In Crassispirinae, up to two sphincters were found within the

buccal tube (Fig. 33 B — Crassispira dysoni, Fig. 33 D— Crassispira

(Crassiclava), and others). Where two sphincters were present, the

anterior was usually much smaller than the posterior. The positions

of the sphincters also vary. A sphincter(s) was defined as anterior in

position, if it lies at a distance of not more than 2.5 marginal tooth

lengths from the mouth opening. The more posterior sphincter of the

buccal tube is shifted backwards in some species, sometimes occu-

pying a position in the mid-proboscis (Fig. 34 A — Antiguraleus

morgani). If the sphincter lies at a distance of more than 2.5 marginal

tooth lengths from the mouth we classified it as intermediate (e.g.

Crassispira dysoni — Fig. 33 B; Crassispira (Striospira) spp. — Fig.

33 F, and others). An intermediate sphincter within the buccal tube

has previously been found only in one species of Splendrillia

(Drilliidae) (Sysoev & Kantor, 1989). Its function is obscure, for it

cannot be used for gripping teeth at the proboscis tip, but may

perhaps be used in transportation of the tooth from the radular sac to

the proboscis tip. The anterior buccal tube sphincter is often absent

and only the intermediate sphincter present (Nquma scalpta,

Naudedrillia praetermissa — Fig. 34 C; Inquisitor spp.; Crassispira

(Striospira) spp. — Fig. 33 F; Miraclathurella bicanalifera). Only

rarely are both sphincters absent and this indicates that separate

marginal teeth are probably not used at the proboscis tip (Burchia

spectabilis — Fig. 33 E, Inquisitor latifasciata — Fig. 33 D).

As in the majority of conoideans, there is in crassispirine species

amore or less well-defined, sac-like enlargement of the buccal tube.

which is lined with a modified epithelium. The structure is associ-

ated with the gripping of single radular teeth at the proboscis tip

(Kantor & Taylor, 1991). From our sections of Crassispirinae, it

appears that the presence and degree of development of the sac-like

enlargement is not correlated with the position of the buccal tube

sphincters (either both anterior and intermediate, or only one may be

present). In three species, namely, C. turricula, C. apicata (Figs 12,

33 D) and C. dysoni (Figs 5, 33 B), the walls of the enlargement were

more muscular than the rest of the buccal tube. No enlargement was

found in Burchia spectabilis (Fig. 33 E), Inquisitor latifasciata (Fig.

34 D), C. tepocana and C. kluthi (Fig. 33 F). In the two former

species, the proboscis tip is able to invert and the separate teeth are

not gripped at the proboscis tip, whilst we have no other information

about C. tepocana and C. kluthi.

An unusual character, previously found only in Splendrillia

(Sysoev & Kantor, 1989), is the presence of an epithelial pad in the

anterior portion of the buccal tube. For some species, marginal teeth

were seen adhering to this pad, probably for more secure fixation.

The epithelial pad was found in several crassispirine species, for

example, Crassispira (Crassiclava) species (Fig. 33 D) and

Crassispira dysoni (Fig. 33 B).

Salivary glands differ greatly in size and histology. Before this

study, only two types of salivary glands were recognised amongst

the Conoidea — acinous and simple tubular (Taylor et al., 1993). The

structure of the glands was thought to be a character useful at

subfamilial level, for instance, differentiating the Mangeliinae and

Raphitominae. However, amongst the crassispirines we were able to

recognise four types of salivary gland. Besides the acinous salivary

glands (found in majority of species) and simple tubular glands

(Haedropleura septangularis — Fig. 34 F; Naudedrillia praetermissa

— Fig. 34 C), we found anastomosing tubular (Figs 33 D, F; 34 C,G)

and glands consisting of a simple tube surrounded by acinous cells

(Funa spp. — Fig. 33 A; Antiguraleus morganus — Fig. 34 A). Some-

times, species attributed to the same genus and even subgenus

possess different types of glands. For example, in Crassispira

(Striospira) tepocana, the glands are anastomosing tubular, while in

C. (S.) kluthi and C.(S.) xanthi they are acinous. Moreover, it should

be emphasised, that there were no correlations between the structure

of the glands and the foregut anatomy. Sometimes in species pos-

sessing the same type of foregut, the glands were of different

histology (eg. Nquma scalpta and Naudedrillia praetermissa — Fig.

34 C; and Crassispira (Crassispira) spp. and Funa spp. — Fig. 33 A).

The Crassispirinae is the only subfamily of the Conoidea, in

which a variety of salivary glands has so far been recorded. It is still

unclear whether the simple tubular salivary glands can be derived

from the acinous type. However, recently, it has been demonstrated

(Ball, Taylor & Andrews, in press) that in the embryonic develop-

ment of Nucella, the salivary ducts are formed first, and the salivary

gland itself appears later at the tip of the duct. From this, it can be

suggested that the simple tubular glands may represent the enlarged

ducts, while the gland itself was not developed. Thus, the simple

tubular salivary glands may originate from the acinous by

paedomorphosis. This may also explain the origin of the anastomo-

sing tubular glands, which could possibly be the result of extensive

coiling of the initial duct.

Shell and radular characters

Radula

Radular morphology has been used extensively to recognise and

classify suprageneric categories within the Conoidea (e.g. Powell,

1966; McLean, 1971; Kilburn, 1988, Taylor er al., 1993). However,

this study has demonstrated that within the Crassispirinae, there is

no great congruence between radular and anatomical characters.

Many crassispirine taxa have quite similar radular teeth (e.g. Figs

4, 11) with the wishbone tooth formed by a robust, pointed, major

element and a smaller, more slender, secondary element. However,

this similarity of radula morphology is not reflected in foregut

anatomy and taxa with the same type of teeth often have widely

different arrangements of the foregut; for example /nquisitor

latifasciata (Figs 17b, 20) and Crassispira harfordiana flucki (Figs

4d, 6). In other cases, some gastropods possess rather similar and

distinctive foregut anatomies, as for example, /nquisitor latifasciata

and Ptychobela suturalis. However, they have very different radular

morphologies, with /. latifasciata having the rather standard crassi-

spirine wishbone form, but P suturalis has the autapomorphic,

awl-shaped, hollow teeth (Fig. 23a).

Although teeth with the wishbone form are found in the sub-

families Turrinae, Clavatulinae and Cochlespirinae, most variation

and the most extreme forms are found within taxa classified as

Crassispirinae. Although rather a disparate range of radular teeth

was found amongst the species we studied, all can probably be

derived from the basic wishbone form. Thus, the derivation of the

rather extreme forms of wishbone teeth seen in Hindsiclava and

Haedropleura can be envisaged by changes in the relative propor-

tion of the elements. In the large, paddle-shaped teeth of Funa and

Vexitomina, a thin, splint-like secondary limb lies along the shaft of

the main tooth. In the most-derived, harpoon-like teeth of

Cheungbeia, a similar small splint-like secondary limb lies along the

edge of the shaft. Finally, the hollow awl-shaped teeth of Ptychobela

consist of two components and can probably be derived from

flattening and fusing of the two wishbone components.

Congruence of shell characters

Shell characters are also a rather poorly correlated with radular

morphology or characters of foregut anatomy. A good example of

this problem is seen in the genera /nquisitor, Funa, and Ptychobela.

These have rather similar shells but the radulae are quite different,

Inquisitor having the standard crassispirine wishbone form, Funa

has bladed, paddle-shaped teeth and Ptychobela possesses awl-

shaped hollow teeth. The allocation of shells to these genera is

difficult without radular evidence and many species have been rather

arbitrarily assigned to genera (e.g. Wells, 1994).

A striking example of the similarity of shell characters in different

families is seen between the pairs of West American species

Crassispira (Striospira) tepocana (see Keen, 1971 fig. 1701) and

Strictispira ericana (Keen, 1971 fig. 1727) and C. (Striospira) xanti

(Keen, 1971 fig. 1702) with Strictispira stillmani (Keen, 1971 fig.

1728). These pairs of similar gastropods have quite different anato-

mies.The Strictispira species (family Strictispiridae) lack the venom

gland, have the buccal mass located at the proboscis tip and possess

very distinctive radula teeth (Kantor & Taylor, 1994).

Parallel evolution of hypodermic feeding

mechanism

In two genera, the radular teeth are quite different from the rest of the

Crassispirinae, which for the most part consist of variations on the

robust wishbone form. In Cheungbeia species, the teeth are long,

slender and harpoon-like, with distal barbs, whilst in Ptychobela the

teeth are pointed, awl shaped and hollow. Although most conoidean

teeth, including the wishbone types can be used at the proboscis tip

for the penetration of prey, the hollow, barbed teeth are regarded as

the more efficient. Taylor et al. (1993) demonstrated that hollow,

barbed teeth had evolved several times within the Conoidea and

these two taxa may represent further evolutionary pathways to the

hypodermic feeding mechanism. The teeth in Cheungbeia and

Ptychobela although quite different in morphology can both be

derived from the crassispiran wishbone form. Although Cheungbeia

is long and gutter-shaped, a small and very thin accessory limb lies

along one side of the shaft (Fig. 26). A similar reduction in the size

of the accessory limb is seen in Funa and Vexitomina which have

paddle-shaped teeth, with the thin secondary limb lying along the

shaft (Figs 23 b-d). In Ptychobela, each tooth is formed from two

more or less equal parts which are fused along one edge and loosely

enrolled. Neither of these two tooth types can be considered as

precursors of the enrolled barbed teeth found in Conus (Coninae,

sensu Taylor et al. 1993) because their foregut anatomy is different.

Both Cheungbeia and Ptychobela, for example, have the modified

epithelium of the rhynchodeum, the elongated oesophagus, and the

Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR

modified venom gland. None of these features is found in Conus.

Moreover, Conidae have lost the radular membrane and possess a

radular caecum for the storage of teeth prior to use.

Comparison between conoidean subfamilies

Three other subfamilies within the Turridae, the Turrinae,

Clavatulinae and Cochlespirinae possess the wishbone type of radular

teeth and their features should be compared with those of the

Crassispirinae.

Members of the Turrinae have wishbone teeth which differ in

morphology from those of the other taxa. The proximal half of each

tooth is divided like a clothes-peg into two more or less equal units

(Fig. 18), with the limbs not detached. This contrasts with the

wishbone teeth in the other subfamilies which have the limbs

unequal in size and thickness and the secondary limb not in struc-

tural continuity with the major limb. Some species of Turrinae also

have a quadrate central tooth with a spine-like central cusp. Species

of Turrinae also differ in a number of anatomical characters; there is

usually no elongation of the oesophagus, no change in the histology

of the venom gland anterior of the nerve ring and the rhynchodeum

is uniformly glandular along its length (Taylor et al., 1993; Taylor,

1994).

The Clavatulinae have wishbone teeth with a large, bladed, major

limb and the secondary limb inserted into a ‘V’ shaped groove.

Additionally, central teeth are usually present. They also have a

medio-lateral nucleus to the operculum rather than the terminal

position found in the other turrid subfamilies. The buccal mass often

lies within the proboscis, but as in Crassispirinae, the oesophagus is

often elongated between the nerve ring and buccal mass and the

venom gland changes to a ciliated duct anterior to the nerve ring

(Kantor, 1990; Taylor er al., 1993).

The Cochlespirinae, represented by Aforia, Antiplanes, and

Cochlespira, have similar wishbone teeth to the Crassispirinae, with

some species possessing a central tooth, and some having plate-like

lateral teeth (Sysoev & Kantor, 1987, 1988; Kantor & Sysoev,

1991). In Cochlespira and Aforia, the venom gland joins the oesopha-

gus some way to the posterior of the buccal mass, but no details are

available for other taxa.

We have found a great variation in the structure of the foregut in

the Crassispirinae and some of these features are shared with the

Turrinae, Clavatulinae and Cochlespirinae. However, rather few

species have been studied from the latter three groups compared

with the more extensive survey of the Crassispirinae. A reappraisal

of the boundaries and relationships of these four subfamilies is

required, but this cannot be attempted before more anatomical

details are available from a much wider range of taxa.

Systematic conclusions

ANTIGURALEUS and associated genera

On the basis of his study of southern African species, Kilburn (1994)

suggested that two genera, Anacithara and Antiguraleus, which had

previously been referred to the subfamily Mangeliinae (as in Powell,

1966), should be transferred to the Crassispirinae on the basis of

radular characters. We studied one of the South African species,

Antiguraleus morgani and confirm that it has a crassispirine type of

radula (Fig. 28a) and also has many foregut characters consistent

with other members of the Crassispirinae. Additionally, we also

sectioned the eastern Australian species, Guraleus costatus, and this

also has an operculum, with a radula and foregut anatomy of the

FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS

Crassispirinae (Fig. 28b). However, another Australian species that

we sectioned, Antiguraleus howelli (Laseron), possesses a typical

mangeliinae anatomy and Powell (1966, fig. 138) also illustrates a

typical mangeliine radula for Antiguraleus murrheus (Webster, 1906).

Our observations suggest that the mostly Australasian species,

currently assigned to the Guraleus group of genera (Antiguraleus,

Paraguraleus, Guraleus and Neoguraleus) (Powell, 1966) represent

at least two different subfamilies, and the whole complex is in great

need of critical revision.

EPIDIRONA

This genus was assigned to the Crassispirinae by Sysoev (In: Taylor

et al., 1993) on the evidence of the radula of the type species

Epidirona hedleyi Iredale, 1931, which has teeth with the wishbone

form similar to many other species in the subfamily, with a robust,

pointed, major limb and a slender secondary limb (Powell, 1964,

plate 229; Powell, 1966, fig. 33). However, our studies show that

Epidirona gabensis has a radula with the wishbone teeth with the

form typical of the subfamily Turrinae, i.e. awl shaped and bifurcat-

ing in the proximal half (Fig. 18) . Similar teeth were illustrated by

Powell (1966, fig. 32) for Epidirona nodulosa Laseron, 1954. In our

phylogenetic analysis Epidirona gabensis was shown to be more

similar, despite some anatomical differences, to the outgroup

Gemmula deshayesii, rather than to the crassispirine species.

However, the type species, Epidirona hedleyi would seem to be a

crassispirine on the evidence of the radula, but we have no anatomi-

cal information to confirm this.

Our conclusions are that Epidirona gabensis and E. nodulosa

should be classified within the subfamily Turrinae. Epidirona hedleyi,

the type species, is likely to be a crassispirine and reference to the

illustrations in Powell (1964, plate 230) shows that it differs from the

other species on shell characters. A new generic name is necessary

for the Epidirona species which possess the Turrinae-type of radular

teeth, however, anatomical or at least radular studies of the other

species is desirable.

TURRIDRUPA

The systematic position of this genus has been uncertain. Powell

(1966; 1967) referred it to the Turrinae on shell characters, but his

illustration (1967 fig. 300) of the radula of 7: jubata would seem to

be a misinterpretation. Subsequently, Kilburn (1983) transferred

Turridrupa to the Clavinae (= Drilliidae), but later, on the basis of

radular characters (Turridrupa bijubata andcincta) concluded (1988

p. 235) that Turridrupa was ‘a primitive crassispirine clade.’ . Addi-

tionally, one of the species, Turridrupa cerithina, was again on

radular characters, transferred to the genus /nquisitor, (Kilburn,

1988, p. 267). Unfortunately, the anatomy is known only for

Turridrupa bijubata which has a quadrate central tooth and wish-

bone marginals (Kilburn, 1988 fig. 40) which are of the clothes-peg

type similar to those of the Turrinae. Also, our phylogenetic analysis

shows that in anatomical characters, Turridrupa bijubata is more

similar to the outgroup Gemmula (Turrinae) and to Epidirona gab-

ensis than any of the Crassispirinae. We have not studied Turridrupa

cerithina which has a different shell morphology from other species

and may well be a crassispirinan. In conclusion, we think that

Turridrupa should be classified in the Turrinae, with T. cerithina

possibly in the Crassispirinae.

CRASSISPIRA and subgenera

At present. the genus Crassispira is usually divided into eight

subgenera. Our phylogenetic analysis shows that Crassispira

(Crassispira) and C. (Striospira) form a monophyletic clade, but

species of the other subgenera appear at widely separated positions

on the cladogram (Fig. 33) and have different anatomies from

Crassispira s.s. Our results suggest that the subgenera of Crassispira

should be raised to full generic status, with perhaps a case for the

retention of Striospira as a subgenus.

CONCLUSIONS

This detailed study of the anterior alimentary system of one sub-

family of conoideans has revealed an extraordinary diversity of

foregut configuration. It is uncertain whether so much variation

exists in other conoidean groups, for far fewer species have been

studied. However, preliminary evidence suggests that a similar

diversity of foreguts exists in the Raphitominae, Clavatulinae and

Terebridae (Taylor, 1990; Taylor et al., 1993; Sysoev & Kantor,

1995; Kantor & Sysoev, 1996). The anatomy of Conus has usually

been taken as being typical of the whole Conoidea, but its foregut is

in many ways rather underived (basal buccal mass, unmodified

rhynchodeal wall, no oesophageal loop, unmodified anterior venom

gland). In fact, Conus can be regarded as just one of many possible

foregut configurations found within the Conoidea. Virtually all the

organs of the foregut can vary in presence or absence, size and

position. Thus for example, the proboscis may be very long or very

short; the buccal mass may be situated near the tip of the proboscis,

at the base or to the posterior; much of the posterior rhynchodeum

may be able to evert thus forming an extended proboscis, or there

may be several of no sphincters in the buccal tube. Indeed, more

extreme conditions exist in the some species of Raphitominae and

Terebridae, where the venom gland, radula, salivary glands and

proboscis have been lost. This diversity of foregut structure of

conoideans likely reflects considerable variation in feeding behav-

iour and methods of prey capture. Unfortunately, apart from Conus,

few details are available for other conoidean taxa (Miller, 1989;

Taylor et al., 1993).

ACKNOWLEDGEMENTS. Weare grateful to Gary Rosenberg, Dick Kilburn,

Jim McLean, Philippe Bouchet, Ian Loch, Winston Ponder, Graham Oliver

and Marco Oliverio for the generous donation of some of the specimens used

in this study. Alexander Sysoev provided discussion and encouragement.

Brian Morton gave much logistic support for the collection of specimens in

Hong Kong. David Cooper expertly prepared many of the serial sections. Yuri

Kantor is grateful to the Royal Society for the air fare enabling him work in

London.

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