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26NOV1987 

PRESENTED 
GFNEH/U LIB3AFV 



Bulletin of the 

British Museum (Natural History) 



Zoology series Vol 52 1987 



British Museum (Natural History) 
London 1987 



Dates of publication of the parts 

Nol . . . 29 January 1987 

No 2 26 February 1987 

No 3 . . 26 March 1987 

No 4 30 April 1987 

No 5 . . . 28 May 1987 

No 6 25 June 1987 

No 7 30 July 1987 

No 8 27 August 1987 



ISSN 0007-1 498 



Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 



Contents 
Zoology Volume 52 



No 1 Miscellanea 



A revision of the genus Pseudovorticella Foissner & Schiffmann, 1974 
(Ciliophora: Peritrichida). By A. Warren 1 

The taxonomic status of the genera Pontigulasia, Lagenodifflugia and 
Zivkovicia (Rhizopoda: Difflugiidae). By C. G. Ogden . . . 13 

A revision of the foraminiferal genus Adercotryma Loeblich & 
Tappan, with a description of A. wrighti sp. nov. from British waters. 
By P. Bronnimann & J. E. Whittaker 21 

Hermit crabs associated with the bryozoan Hippoporidra in British 

waters. By J. D. D. Bishop 29 

The first zoea of three Pachygrapsus species and of Cataleptodius 
floridanus (Gibbes) from Bermuda and Mediterranean (Crustacea: 
Decapoda: Brachyura). By R. W. Ingle 31 

A classification of the phylum Sipuncula. By P. E. Gibbs & E. B. 

Cutler 43 

Two new species ofGarra (Teleostei: Cyprinidae) from the Arabian 
peninsula. By K. E. Banister 59 

No 2 A revision of the Suctoria (Ciliophora, Kinetofragminophora) 5. The 

Paracineta and Corynophora problem. By Colin R. Curds . . . 71 

No 3 Notes on spiders of the family Salticidae 1 . The genera Spartaeus, 

MintoniaandTaraxella.RyF.R.'Wan\ess 107 

No 4 Mites of the genus Holoparasitus Oudemans, 1936 (Mesostigmata: 

Parasitidae) in the British Isles. By K. H. Hyatt 139 

No 5 The phylogenetic position of the Yugoslavian cyprinid fish genus 

Aulopyge Heckel, 1841, with an appraisal of the genus Barbus Cuvier 

& Cloquet, 1816 and the subfamily Cyprininae. By Gordon J. Howes . 1 65 

No 6 Revision of the genera Acineria, Trimyema, and Trochiliopsis 

(Protozoa, Ciliophora). By H. Augustin, W. Foissner & H. Adam . 197 

No 7 The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) 
with a systematic review, a synopsis of Pipistrellus and Eptesicus, and 
the descriptions of a new genus and subgenus. By J. E. Hill & D. L. 
Harrison . 225 

No 8 Notes on some species of the genus Amathia (Bryozoa, 

Ctenostomata). By P. J. Chimonides 307 






Bulletin of the 

British Museum (Natural History) 



Miscellanea 



Zoology series Vol52 No 1 29 January 1987 



The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four 
scientific series. Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and 
an Historical series. 

Papers in the Bulletin are primarily the results of research carried out on the unique and 
ever-crowing collections of the Museum, both by the scientific staff of the Museum and by 
specialists from elsewhere who make use of the Museum's resources. Many of the papers are 
works of reference that will remain indispensable for years to come. 

Parts are published at irregular intervals as they become ready, each is complete in itself, 
available separately, and individually priced. Volumes contain about 300 pages and several 
volumes may appear within a calendar year. Subscriptions may be placed for one or more of 
the series on either an Annual or Per Volume basis. Prices vary according to the contents of 
the individual parts. Orders and enquiries should be sent to: 



Publications Sales, 

British Museum (Natural History), 
Cromwell Road, 

London SW7 5BD, 
England. 



World List abbreviation: Bull. Br. Mus. nat. Hist. (Zool.) 



Trustees of the British Museum (Natural History), 1986 



The Zoology Series is edited in the Museum's Department of Zoology 

Keeper of Zoology : Mr J. F. Peake 
Editor of Bulletin : Dr C. R. Curds 
Assistant Editor Mr C. G. Ogden 



ISBN 0565 05025 7 

ISSN 0007- 1498 Zo ?l, g l T Se T , 

Vol 52 No. 1 pp 1-70 

British Museum (Natural History) 

Cromwell Road , 

London SW7 5BD Issued 29 January 1987 






Miscellanea 




Contents 



A revision of the genus Pseudovorticella Foissner & Schiffmann, 1974 (Ciliophora: 
Peritrichida). By A. Warren 

The taxonomic status of the genera Pontigulasia, Lagenodifflugia and Zivkovicia 
(Rhizopoda: Difflugiidae). By C. G. Ogden 

A revision of the foraminiferal genus Adercotryma Loeblich & Tappan, with a description 
of A. wrighti sp. nov. from British waters. By P. Bronnimann & J. E. Whittaker . 

Hermit crabs associated with the bryozoan Hippoporidra in British waters. By J. D. D. 
Bishop 

The first zoea of three Pachygrapsus species and of Cataleptodius floridanus (Gibbes) from 
Bermuda and Mediterranean (Crustacea: Decapoda: Brachyura). By R. W. Ingle. 

A classification of the phylum Sipuncula. By P. E. Gibbs & E. B. Cutler .... 



Page 
1 

13 
21 
29 

31 

43 



Two new species of Garra (Teleostei: Cyprinidae) from the Arabian peninsula. By 
K. E. Banister 



59 



A revision of the genus Pseudovorticella Foissner & 
Schiffmann, 1974 (Ciliophora: Peritrichida) 

A. Warren 

Department of Zoology, British Museum (Natural History), Cromwell Road, 



London SW7 5BD 



Introduction 

The genus Pseudovorticella was erected by Foissner & Schiffmann ( 1 974) to include those peritrichs 
which are morphologically similar to Vorticella but which have a reticulate silver line system 
with lines running vertically as well as horizontally. The reticulate pattern of silver lines under- 
lies a system of pellicular tubercles which covers the entire zooid surface except the disc and 
infundibulum. 

Pellicular tubercles have been studied by several workers over the past century. Schroder (1906) 
showed that the tubercles of Pseudovorticella monilata are surface features, the distribution 
of which corresponds to that of the underlying striations. Ultrastructural studies by TEM 
(Kawamura, 1 973) and SEM (Carey & Warren, 1 983) have confirmed this observation. Kawamura 
(1973) also showed that each tubercle of P. monilata is a semisphere, about 2-0 um in diameter, and 
contains a sphere of electron dense material. Further investigations using histochemical staining 
(Faure-Fremiet&Thaureaux, 1 944; Pratt & Rosen, 1983) and microanalysis (Pratt & Rosen, 1983) 
indicate that the tubercles contain paraglycogen. The function of the tubercles is not known 
although it has been suggested that they may aid predator avoidance (Spoon, 1975). 

Foissner & Schiffmann (1974) noted that the silver line system is particularly useful for species 
diagnosis in Pseudovorticella, and biometric analyses have been carried out on several species 
(Foissner & Schiffmann, 1974 & 1975; Foissner, 1979). Parameters which are of particular taxo- 
nomic value include the total number of silver lines per zooid and dimensions of the grids formed 
by the intersecting vertical and horizontal lines. Morphological features traditionally used in 
vorticellid taxonomy are also useful diagnostic characters for the species of Pseudovorticella; these 
include the size and shape of the zooid, the number and position(s) of the contractile vacuole(s) and 
the shape and position of the macronucleus (Noland & Finley, 1931; Foissner, 1979; Warren, 
1986). 

Sixteen species of Pseudovorticella are recognised, twelve of which originally belonged to the 
genus Vorticella. A key to their identification is provided. 



Systematics 

In the scheme adopted by the Committee on Systematics and Evolution of the Society of Proto- 
zoologists (Levine et al., 1980), the taxonomic position of the genus Pseudovorticella was given as 
follows: 

Phylum: Ciliophora Doflein, 1 90 1 

Class: Oligohymenophora de Puytorac et al., 1974 

Subclass: Peri trichia Stein, 1859 

Order: Peritrichida Stein, 1859 

Suborder: Sessilina Kahl, 1933 

Family: Vorticellidae Ehrenberg, 1838 

Genus: Pseudovorticella Foissner & Schiffmann, 1974 



Bull. Br. Mus. not. Hist. (Zool.) 52(1): 1-12 Issued 29 January 1987 



2 A. WARREN 

Diagnosis 

Solitary bell-shaped zooids borne upon a spirally contractile stalk. In all respects save one, the 
body and stalk of Pseudovorticella resemble those of Vorticella from which it cannot be differen- 
tiated until impregnated with silver, which reveals a reticulate silver line pattern quite unlike 
that of Vorticella (see Warren, 1986). In addition to Vorticella this genus could be mistaken for 
Haplocaulis in which the stalk contracts in a zigzag rather than a helical manner. 



Key to the species of Pseudovorticella 

1 With endosymbiotic zoochlorellae 2 

Without endosymbiotic zoochlorellae 3 

2 Zooid about 40 um long; macronucleus C-shaped ... P. zooanthelligera (Fig. 9b) 
Zooid 75-95 um long; macronucleus J-shaped P. chlorelligera (Fig. la) 

3 Diameter of peristomial lip less than or equal to maximum body width 4 

Diameter of peristomial lip greater than maximum body width 10 

4 Diameter of peristomial lip less than maximum body width 5 

Diameter of peristomial lip equal to maximum body width 6 

5 Macronucleus J-shaped P. difficilis (Fig. 1 c & d) 

Macronucleus C-shaped P. papillata (Fig. 5c) 

6 Body length less than x 2 maximum body width 7 

Body length at least x 2 maximum body width P. micata (Fig. 2b) 

7 One contractile vacuole 8 

Two contractile vacuoles P. sphagni (Fig. 7b) 

8 Macronucleus lies vertical with respect to major axis of zooid 9 

Macronucleus lies horizontal with respect to major axis of zooid . . P. stilleri (Fig. 9a) 

9 Zooid 65-80 urn long and with 44-54 transverse striations . . . .P. quadrat a (Fig. 7a) 
Zooid 35-45 urn long and with 20-33 transverse striations . . . P. sauwaldensis (Fig. 8) 

10 Zooid with two contractile vacuoles 11 

Zooid with one contractile vacuole 13 

1 1 Zooid with centrally located constriction; scopular region rounded . P. margaritata (Fig. 2a) 
Zooid without centrally located constriction; scopular region tapers towards stalk . . .12 

12 Zooid 40-45 urn long; stalk length x 16- 18 zooid length .... P. mollis (Fig. 3a) 
Zooid 60-70 um long; stalk x 3 zooid length P. monilata (Fig. 3b, c & d) 

13 Diameter of peristomial lip less than body length 14 

Diameter of peristomial lip greater than body length P. punctata (Fig. 6) 

14 Zooid less than x 3 maximum body width 15 

Zooid length greater than x 3 maximum body width .... P. mutans (Fig. 4 c & d) 

1 5 Zooid 50-70 urn long x 22^48 um wide; typically marine . . .P. nebulifera (Fig. 4 a & b) 
Zooid 32-50 um long x 20 um wide; typically freshwater . P. pseudocampanula (Fig. 5 a & b) 



Description of Species 

P. chlorelligera (Kahl, 1935) Jankowski, 1976 

V. margaritata f. chlorelligera Kahl, 1935 

P. margaritata f. chlorelligera (Kahl, 1935) Foissner & Schiffmann, 1975 

DIAGNOSIS (Fig. la & b). Zooid inverted bell-shaped, 78-95 um longx 50 um wide; peristomial lip 80 um 
diameter; infundibulum reaches half body length; macronucleus J-shaped; numerous endosymbiotic zoo- 
chlorellae present in cytoplasm; zooid has a total of 33-53 (mean 47-7) transverse striations; grid size 



GENUS PSEUDOVORTICELLA 





Fig. 1. (a) P. chlorelligera zooid, bar 50 um; (b) telotroch, bar = 25 um (after Foissner & Schiffmann, 
1975); (c) P. difficilis (after Kahl, 1935); (d) P. difficilis, bar = 50um (after Foissner & Schiffmann, 
1975; called P. difficilis var. magnistriata). 



1-3-3-2 um x 2-0-4-5 um; zooid surface with 15-28 (mean 22) pellicular pores per 100 um 2 ; telotroch nearly 
cylindrical in shape and with a prominent epistomial membrane (Fig. lb). 

HABITAT. Freshwater. 

P. difficilis (Kah\, 1933) Jankowski, 1976 
V. difficilis Kahl, 1933 
P. difficilis var. magnistriata Foissner & Schiffmann, 1974 

DIAGNOSIS (Fig. Ic & d). Zooid 60-140 um long x 40-70 um wide; diameter of peristomial lip less than 
maximum body width; infundibulum reaches half body length; single contractile vacuole situated in upper 
part of zooid close to infundibulum; macronucleus J-shaped; 39-49 (mean 43-9) transverse striations per 
zooid; grid size 3- 1-4-7 um x 2-7-3-4 jam; spasmoneme with numerous thecoplasmic granules. 

HABITAT. Freshwater or marine. 

P. margaritata (Fromentel, 1874) Jankowski, 1976 

DIAGNOSIS (Fig. 2a). Zooid inverted bell-shaped, 59-70 um long x 50 um wide, with a slight constriction in the 
central region and rounded at the scopular end; peristomial lip 70 um in diameter; two contractile vacuoles 
situated in anterior part of zooid; macronucleus C-shaped and situated in centre of zooid. 

HABITAT. Freshwater, particularly eutrophic lakes and stagnant water. 
REMARKS. This species has been redescribed by Kahl (1935) and Stiller (1971). 

P. micata (Kahl, 1933) nov. comb. 
V.micata Kahl, 1933 

DIAGNOSIS (Fig. 2b). Zooid elongate, 65 um long x 25 um wide; peristomial lip 25 um in diameter; disc flat and 
slightly elevated above peristome; infundibulum reaches one third zooid length; contractile vacuole situated 
in upper part of zooid close to infundibulum. 

HABITAT. Marine. 



A. WARREN 





Fig. 2. (a) P. margaritata , bar = 25 urn (composite from Kahl, 1935 and Stiller, 1971); (b) P. micata, 

bar = 25 um (after Kahl, 1935). 



P. mollis (Stokes, 1887) nov. comb. 
V. mollis Stokes, 1887 

DIAGNOSIS (Fig. 3a). Zooid inverted bell-shaped, 40-45 urn long x 25 um wide; peristomial lip 40 um in 
diameter; infundibulum reaches one third body length; two contractile vacuoles situated in anterior part of 
zooid; stalk x 16-1 8 zooid length. 

HABITAT. Freshwater 

REMARKS. Although this species was not drawn by Stokes (1887), it has been observed and figured by 
Nenninger(1948). 

P. monilata (Tatem, 1870) Foissner & Schiffmann, 1974 

V. lockwoodii Stokes 1884 
V. monilata Tatem, 1870 

DIAGNOSIS (Fig. 3b, c & d). Zooid inverted bell-shaped, 45-70 urn long x 40-^45 urn wide; peristomial lip 50 um 
in diameter; infundibulum reaches half body length; two contractile vacuoles situated in anterior part of 
zooid; macronucleus J-shaped; 31-41 (mean 35-3) transverse striations per zooid; grid size 2-5-3-5 umx 
1 -5-2-5 um; stalk x3 body length; spasmoneme with thecoplasmic granules; telotroch cone-shaped with 
prominent epistomial membrane. 

HABITAT. Freshwater, often forming pseudocolonies; Pratt & Rosen (1983) reported large numbers of 
Pseudovorticella (Vorticella) monilata attached the Cyanobacterium Anabaenaflos-aquae. 

P. mutatis (Penard, 1922) Foissner, 1979 
V. mutans Penard, 1922 

DIAGNOSIS (Fig. 4c & d). Zooid inverted bell-shaped, 65-95 um long x 1 8-25 um wide; peristomial lip 25 um in 
diameter; disc convex; infundibulum reaches half body length; contractile vacuole situated in upper half of 
body close to infundibulum; macronucleus J-shaped; zooid has 40-^47 (mean 43) transverse striations; grid 



GENUS PSEUDOVORTICELLA 





Fig. 3. (a) P. mollis, bar = 25 ^m (after Nenninger, 1948); (b) P. monilata showing oral ciliation (detail 
from Patsch, 1974); (c) zooid, bar = 25 urn; (d) telotroch (after Foissner, 1979). G = germinal kinety; 
H = haplokinety; P l , P 2 , P 3 = 1 , 2, 3, peniculus; PO = polykinety. 





Fig. 4. (a) P. nebulifera zooid, bar = 25 ^m (after Noland & Finley, 1 93 1 ); (b) telotroch (after Barlow & 
Finley, 1976b); (c)P. mutans telotroch; (d) zooid, bar = 25 urn (after Foissner, 1979). 



6 A. WARREN 

size 1 -4-1 -5 umx 1-5-2-2 um; stalk x5 body length and 8-0 urn wide; spasmoneme with thecoplasmic 
granules; telotroch with prominent epistomial membrane. 

HABITAT. Freshwater. 

P. nebulifera (Miiller, 1786) Jankowski, 1976 
V. nebulifera Miiller, 1*786 

DIAGNOSIS (Fig. 4a & b). Zooid inverted bell-shaped, 38-78 urn (mean 60 urn) long x 22-48 um (mean 37 um) 
wide; slightly constricted beneath peristomial lip which measures 32-66 um (mean 53 um) in diameter; 
single contractile vacuole situated close to infundibulum; macronucleus J-shaped; stalk 50-800 um (mean 
1 50 um) long x 3-5-6-0 um (mean 4-7 um) wide; spasmoneme with thecoplasmic granules; telotroch 47-75 um 
(mean 60 um) long; cyst 37 um in diameter. 

HABITAT. Marine or freshwater. 

REMARKS. Redescribed by Noland & Finley (1931); for telotroch and SEM studies, see Barlow & Finley 
(1976a&b). 

P.papillata (Stiller) Jankowski, 1976 
V. microstoma f. monilata Stiller (see Stiller, 1971) 

DIAGNOSIS (Fig. 5c). Zooid 35-80 um (mean 55 jam) long x 22-50 um (mean 35 um) wide, the maximum 
body width being the mid region of the zooid; peristomial lip 12-25 um (mean 23 urn) in diameter; disc 
convex; infundibulum reaches one third body length; contractile vacuole situated in anterior part of zooid; 
macronucleus C-shaped and lies longitudinally with respect to major axis of zooid. 

HABITAT. Freshwater, particularly under conditions of high biochemical oxygen demand (BOD 5 ). 





Fig. 5. (a) P. pseudocampanula relaxed zooid, bar = 25 um; (b) contracted zooid (after Foissner, 1 979); 
(c) P.papillata, bar = 25 um (after Stiller, 1971). 



GENUS PSEUDOVORTICELLA 




Fig. 6. P. punctata, (a) bar = 25 urn (after Dons, 1918); (b) after Stiller (1946) (called Vorticella 
subconica); (c) after Kahl (1935) (called Vorticella perlata). 



P. pseudocampanula Foissner, 1 979 

DIAGNOSIS (Fig. 5a & b). Zooid conical/inverted bell-shaped, 32-50 um (mean 40 um) long x 20 urn wide; 
peristomial lip 35 um in diameter; upon contraction, peristomial lip becomes puckered (Fig. 5b); infundibu- 
lum reaches half body length; contractile vacuole situated close to infundibulum; macronucleus J-shaped; 
zooid has 44-51 (mean 46-6) transverse striations; grid size 1-3-2-6 urn x 1-5-3-0 urn; stalk x 7 body length; 
thecoplasmic granules present on spasmoneme. 

HABITAT. Freshwater. 

P. punctata (Dons 1918) nov. comb. 

V. punctata Dons, 1918 

V. subconica Stiller, 1946 

P. subconica (Stiller, 1946) Jankowski, 1976 

DIAGNOSIS (Fig. 6). Zooid conical or inverted bell-shaped, 40-50 um Iongx40um wide; peristomial lip 
50-55 um in diameter; disc convex; infundibulum reaches one third body length; contractile vacuole situated 
in upper part of zooid; macronucleus J-shaped; stalk x 4-5 body length and 4-0 um wide. 

HABITAT. Marine. 

P. quadrata Foissner, 1 979 

DIAGNOSIS (Fig. 7a). Zooid 65-80 um (mean 70 um) long x 55 um wide; peristomial lip 60 um in diameter; 
infundibulum reaches half body length; contractile vacuole situated in anterior part of zooid; macronucleus 
J-shaped; zooid has 44-54 (mean 48-3) transverse striations; grid size 1-5-2-8 um x 1-3 x 2-7 um; stalk x 7 
body length and 9-0 um wide; spasmoneme with thecoplasmic granules. 

HABITAT. Freshwater. 

P. sauwaldensis Foissner & Schiffmann, 1979 

DIAGNOSIS (Fig. 8). Zooid shape variable, usually inverted bell-shaped 35^5 um long x 20 um wide; 
peristomial lip 20 um in diameter and 3-0 um thick; disc convex; infundibulum reaches half body length; 



A. WARREN 





a b 

Fig. 7. (a) P. quadrata, bar = 25 nm; (b) P. sphagni, bar = 25 um (after Foissner, 1 979). 





Fig. 8. P. sauwaldensis (a) normal zooid, bar = 20 um; (b) contracted zooid; (c) showing variability of 
macronucleus and zooid shape, bar = 20 urn (after Foissner & Schiffmann, 1979). 

contractile vacuole situated in upper part of zooid close to infundibulum; macronucleus vermiform, variable 
in shape and situated longitudinally with respect to major body axis; pellicle has 20-33 (mean 29) transverse 
striations; grid size 0-9-1-5 um x 0-7-2-5 um; stalk x 1-3 body length. 

HABITAT. Freshwater. 



GENUS PSEUDOVORTICELLA 




Fig. 9. (a) P. stilleri, bar = 50 um (after Stiller, 1 963); (b) P. zooanthelligera, bar = 25 urn (after Stiller, 

1968). EZ = endosymbiotic zoochlorellae. 



P. sphagni Foissner, 1979 

DIAGNOSIS (Fig. 7b). Zooid inverted bell-shaped, 40-50 um long x 30 um wide; peristomial lip 30 um in 
diameter; infundibulum reaches one third body length; two contractile vacuoles situated in anterior part of 
zooid; macronucleus J-shaped with elongate distal arm; zooid has 34-37 (mean 35-5) transverse striations; 
grid size 1 -6-1 -9 um x 2-5-2-7 um. 

HABITAT. Freshwater, originally isolated from Sphagnum bogs. 

P. stilleri n. sp. 

V. campanula f. monilata Stiller, 1963 

DIAGNOSIS (Fig. 9a). Zooid inverted bell-shaped, 85 um long x 80 um wide; peristomial lip 80 um in diameter; 
infundibulum reaches half body length; macronucleus C-shaped and lies horizontally across centre of zooid. 

HABITAT. Freshwater, attached to the duckweed Lemna minor. 

P. zooanthelligera (Stiller, 1968) nov. comb. 
V. zooanthelligera Stiller, 1968 

DIAGNOSIS (Fig. 9b). Zooid inverted bell-shaped, 40-42 um long x 40 jam wide; peristomial lip 50 um 
in diameter; disc flat; infundibulum reaches one third body length; macronucleus C-shaped and lies 
longitudinally in zooid; cytoplasm contains numerous endosymbiotic zoochlorellae; stalk x 5 body length. 

HABITAT. Freshwater. 



Incertae sedis 

Pseudovorticella sp. (Graham & -Graham, 1978) nov. comb. 
Vorticella sp. Graham & Graham, 1978 

Graham & Graham (1978) made an ultrastructural study of a vorticellid (Vorticella sp.) furnished with 
pellicular tubercles and containing endosymbiotic zoochlorellae. The presence of pellicular tubercles suggests 



10 A. WARREN 

that this organism should belong to the genus Pseudovorticella. However other important diagnostic features, 
for example the macronucleus, contractile vacuole(s) and shape of the relaxed zooid, were not recorded. Only 
when such data is available will it be possible to determine the exact status of this organism. 



Acknowledgements 

I would like to thank Dr W. Foissner for his helpful criticism of the manuscript. 

References 

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Levine, N. D., Corliss, J. O., Cox, F. E. G., Deroux, G., Grain, J., Honigberg, B. M., Leedale, G. F., Loeblich, 

A. R., Lorn, J., Lynn, D., Merinfield, E. G., Page, F. C., Poljansky, G., Sprague , V., Vavra, J. & Wallace, 

F. G. 1980. A newly revised classification of the Protozoa. Journal of Protozoology 27: 37-58. 
Miiller, O. F. 1786. Animalcula Infusoria Fluviatilia et Marina. Havniae et Lipsiae. 367 pp. 
Nenninger, U. 1948. Die Peritrichen der Umgebung von Erlangen mit besonderer Beriicksichtigung ihrer 

Wirtsspezifitat. Zoologische Jahrbiicher (Systematik) 77: 169-266. 
Noland, L. E. & Finley, H. E. 1931. Studies on the taxonomy of the genus Vorticella. Transactions of the 

American Microscopical Society 50: 81-125. 
Patsch, B. 1974. Die Aufwuchsciliaten des Naturlehrparks haus Wildenrath. Monographische Bearbeitung 

der Morphologic und Okologie. Arbeiten aus dem Institutfur Landwirtschaftliche Zoologie und Bienkunde 

No. 1.1-78. 
Penard, E. 1922. Etudes sur les Infusoires d'Eau Douce. 331 pp. George & Cie, Geneva. 



GENUS PSEUDOVORTICELLA 1 1 

Pratt, J. R. & Rosen, B. H. 1983. Association of species of Vorticella (Peritrichida) and planktonic algae. 
Transactions of the American Microscopical Society 102: 48-54. 

Puytorac, P. de, Batisse, A., Bohatier, J., Corliss, J. O., Deroux, G., Didier, P., Dragesco, J., Fryd-Versavel, 
G., Grain, J., Groliere, C. A., Hovasse, R., Itfode, F., Laval, M., Roque, M., Savoie, A. & Tuffrau, M. 1974. 
Proposition d'une classification du phylum Ciliophora Doflein, 1901 (Reunion de Systematique, 
Clermont-Ferrand). Compte Rendu Hebdomadaire des Seances de I'Academie des Sciences. Paris 278: 
2799-2802. 

Schroder, O. 1906. Beitrage zur Kenntnis von Vorticella monilata Tatem. Archiv fur Protistenkunde 7: 
395^10. 

Sondheim, M. 1929. Protozoen aus der Voeltzkowschen Reisen in Madagaskar und Ostafrika. Abhandlungen 
hrsg. von der Senckenbergischen Naturforschenden Gesellschaft 41: 285-313. 

Spoon, D. M. 1975. Survey, Ecology and Systematics of the Upper Potomac Estuary Biota; Aufwuchs Micro- 
fauna, Phase 1. Final Report. Water Resources Center, Washington Technical Institute, Washington. 117 
pp. 

Stein, F. 1859. Der Organismus der Infusionsthiere nach eingenen Forschungen in Systematischer Reihenfolge 
bearbeitet I. 206 pp. Leipzig. 

Stiller, J. 1940. Beitrage zur Peritrichenfauna des Groben Ploner Sees in Holstein. Archiv fur Hydrobiologie 
38: 263-285. 

1 946. Beitrage zur Kenntnis der Peritrichenfauna der Adria bei Split. Annales Historico-Naturales Musei 

Nationalis Hungarici 39: 59-74. 

1963. Zur Limnologie der Natrongewasser Ungarns. I Der Natronsee Nagyszek und seine Peritrichen- 
fauna. Internationale Revue der Gesamten Hydrobiologie und Hydrographie 48: 603-612. 

1968. Peritriche Ciliaten Okologisch verschiedener Biotope von Rovinj und Umgebung. Acta Zoologica 

Academiae Scientiarum Hungaricae 14: 185-21 1. 

1971. Szajoszorus Csillosok-Peritricha. Fauna Hungaricae 105: 1-245. 



Stokes, A. C. 1883. A new vorticellid. American Monthly Microscopical Journal 4: 208. 

1884. A new infusorien belonging to the genus Vorticella. American Naturalist 18: 829-830. 

1 887. Notices of new fresh water infusoria. Proceedings of the American Philosophical Society 24: 

244-255. 
Tatem, J. G. 1870. A contribution to the teratology of the infusoria. Monthly Microscopical Journal 3: 

194-195. 
Warren, A. 1986. A revision of the genus Vorticella (Ciliophora: Peritrichida). Bulletin of the British Museum 

(Natural History). Zoology Series 50(1): 1-57. 

Manuscript accepted for publication 5 January 1986 



Index 

Index of extant species; annotated list of nominal species. 

P. anabaenae (Stiller, 1940) Jankowski, 1976 was transferred to the genus Haptocaulis by Stiller (1971). 

P. Morelligera(Kah\, 1935) Jankowski, 1976 2 

P.chlamydophora(PemLTd, 1922) Jankowski, 1976= Vorticella vestita Stokes, 1 883 (see Warren, 1986). 

P. difficilis (Kahl, 1933) Jankowski, 1976 3 

P. difficilis var. magnistriata Foissner & Schiffmann, 1974 = P. difficilis. 

P. lima (Kahl, 1935) Jankowski, 1976. This species appears to have pellicular granules rather than 

tubercles; it should therefore remain in the genus Vorticella (V. lima) until it has been redescribed. 

P. margaritata (Fromenlel, 1874) Jankowski, 1976 3 

P. micata (Kahl, 1933) nov. comb 3 

P. mollis (Stokes, 1887) nov. comb 4 

P. monilata (Tatem, 1870) Foissner & Schiffmann, 1974 4 

P. wwtaAW (Penard, 1922) Jankowski, 1976 4 

P. nebulif era (Mutter, 1786) Jankowski, 1976 6 

P. papillata (Stiller) Jankowski, 1976 6 

P. pelagica (Gajewskaja, 1933) Jankowski, 1976 was transferred to the genus Haplocaulis by Stiller 

(1971). 



1 2 A. WARREN 

P. perlata (Kahl, 1933) Jankowski, 1916 = P. punctata. 

P. plicata (Gourret & Roeser, 1886) Jankowski, 1976 appears to be identical to Vorticella elongata 

Fromentel, 1874 (Warren, 1986). 

P. pseudocampanula Foissner, \979 7 

P. punctata (Dons, 1918) nov. comb 7 

P. quadrat a Foissner, 1979 7 

P. sauwaldensis Foissner &Schiffmann, 1979 7 

P. sphagni Foissner, 1979 9 

P. stilleri (Stiller, 1963) n. sp 9 

P. subconica (Stiller, 1946) Jankowski, 1976 appears to be identical to P. punctata. 

P. vestita (Stokes, 1883) Jankowski, 1976. A membranous alveolar covering overlays the pellicle of this 

species. There is, however, no evidence of pellicular tubercles or of an underlying reticulate pattern of 

silver lines. This species should therefore remain in the genus Vorticella ( V. vestita) until it has been 

redescribed. 
P. voeltzkowi (Sondheim, 1929) Jankowski, 1976. This species has spine-like projections on its pellicle. 

There is, however, no evidence that it has either pellicular tubercles or a reticulate pattern of silver 

lines. It should therefore remain in the genus Vorticella (V. voeltzkowi) until a redescription is 

available. 
P. zooanthelligera (Stiller, 1968) nov. comb 9 



The taxonomic status of the genera Pontigulasia, 
Lagenodifflugia and Zivkovida (Rhi/opoda: 
Difflugiidae) 

Colin G. Ogden 

Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD 

Introduction 

The significance of the different structural elements utilised to provide an internal dividing wall in 
the shell of the genus Pontigulasia Rhumbler, 1896 was employed by Ogden (1983) to divide the 
genus into two, with the creation of a new genus Zivkovida Ogden, 1983. At the same time a new 
genus Lagenodifflugia was erected by Medioli & Scott (1983) to accommodate Difflugia vas Leidy, 
1 874, a species that he (Leidy, 1 879) later redescribed as a variety of Difflugia pyr if ormis. Medioli & 
Scott (1983) considered their new genus to be distinct from species of Difflugia because the shell was 
divided into a bulbous main part and a neck, the two parts being separated by an internal 
diaphragm pierced by a single, central, usually large orifice. In the same work Medioli & Scott 
redefined the genus Pontigulasia, describing the main characters as a constriction of the neck 
marking the position where a perforated internal diaphragm often extends across this region, and 
at times a bent neck combined with the internal restriction simulating a spiral arrangement. They 
also stated that the diaphragm is seldom present in fossilised forms. The amendment of the 
diagnosis to contain a 'bent neck' was made to allow Medioli & Scott to include some 
'Lecquereusia-like' specimens in their description of Pontigulasia compressa, and to suggest the 
possibility of combining the genera Lesquereusia and Pontigulasia, the former being the senior 
synonym. 

I am most grateful to Dr Drew Haman and Dr Georges Merinfeld for directing my attention to 
the nomenclature problems posed by the creation of these two new genera. The present report is an 
attempt to clarify the status and diagnosis of the three genera Pontigulasia, Lagenodifflugia and 
Zivkovida. 



Taxonomy 

The taxonomic problem caused by the creation of two new genera relates to the interpretation and 
validity of the structures and openings found associated with the inner dividing wall. There are 
only a few descriptions and figures of these features around which the diagnoses have been erected. 
One reason for this is the difficulty of trying to see inside the shell either en face or laterally due 
mainly to the opacity of the mineral particles of which it is constructed. Several techniques have 
been tried to overcome this, for example immersion in clove oil or Canada balsam and deminerali- 
sation by hydrofluoric acid. Modern techniques have now obviated this problem and allow a new 
appraisal of this feature (Ogden, 1983). 

Each genus will be reassessed here on the basis of the available descriptions and the author's 
earlier observations. 

Pontigulasia Rhumbler, 1 896 

The first author to recognise the inner dividing wall was Rhumbler (1896) who erected the genus 
Pontigulasia using this feature as a major diagnostic character to separate it from Difflugia, and 
described and clearly figured it as an internal 'schlundbriicke'-throat bridge. He described three 



Bull. Br. Mus. nat. Hist. (Zool.) 52(1): 13-17 Issued 29 January 1987 



14 C. G. OGDEN 

new species belonging to this genus, P. compressa, P. incisa and P. spiralis. Since then Hopkinson 
(1919) has redescribed P. compressa under a new name P. r humbler i, the name compressa was 
preoccupied by Carter's (1864) species, and P. incisa has become a synonym off. elisa (Penard, 
1 893), both the earlier descriptions being for species of Difflugia. The authoritative date for P. 
rhumbleri is considered to be Hopkinson's redescription (1919), not the note mentioned as an 
addition to P. elisa on p. 162 of Cash & Hopkinson (1909) where Hopkinson suggests that 
'Rhumbler's P. compressa (which might now be called P. Rhumbleri}\ his brackets. Furthermore, 
the date of Rhumbler's work is erroneously quoted by Cash et al. (1919) and Loeblich & Tappan 
(1964) as 1895, which is the date of presentation, whereas the publication date of the volume was 
1896, as listed in Penard's (1902) bibliography. 

The internal bridge has been redescribed by Ogden ( 1 983), who examined the dividing wall in situ 
using scanning electron microscopy, and showed it to be correctly termed a bridge, as it represented 
a rather weak connection between the two lateral walls of the compressed shell in P. rhumbleri. It is 
often difficult to see by optical microscopy because the shells are laterally compressed, and in this 
the normal viewing position the bridge appears to be a dark floating band situated centrally in the 
neck region without any apparent connection to the shell wall. This is not surprising, because each 
junction of the bridge with the wall is about a sixth of the shell diameter at this point. 

Although Rhumbler described three species, since that time two of these P. incisa and P. spiralis 
have been considered to be synonyms (Cash & Hopkinson, 1909), and it was not until Leoblich & 
Tappan (1964) that a type species, P. rhumbleri, was designated. 

DIAGNOSIS. Shell pyriform, sometimes with a constriction of the neck, either circular or compressed 
in transverse section; composed mainly of agglutinate mineral particles with some diatom frustules 
or siliceous plates, bound by a network of organic cement; aperture terminal, circular; internally 
the shell is divided into two regions by a narrow bridge, made mainly of organic cement with some 
agglutinate particles, stretched between the lateral walls at about one third of the body length from 
the aperture. Type species P. rhumbleri Hopkinson, 1919, with four other species (Chardez, 1985), 
P. compressoidea Jung, 1 942; P. elisa (Penard, 1 893); P. sarrazinensis Chardez & Caspar, 1 984 and 
P. spiralis Rhumbler, 1896. 

Lagenodifflugia Medioli & Scott, 1983 

This genus was established for the single species Difflugia vas Leidy, 1874. After his original 
description Leidy later considered (1879) that this species was a variety of Difflugia pyriformis, 
from which it differed by a constriction of the neck, there was no reference to an internal structure 
associated with the constriction. Penard (1902) transferred this species to Pontigulasia and con- 
sidered it to be a synonym of his new species P. spectabilis. It was corrected to the valid binomen P. 
vas by Schoutenden (1906), and spectabilis has since been considered a synonym of vas. As P. vas 
was described as having a similar external constriction to spectabilis, the assumption was that it 
had two openings bisecting the internal diaphragm. Notwithstanding this, Stump (1935, 1936 and 
1 943) in a series of experiments with specimens he initially thought were Difflugia oblonga, later 
found that in sectioned shells there was an internal division with a single opening and subsequently 
described them as Pontigulasia vas. Although Stump does not clearly state that his specimens had a 
single, central opening, his diagrams without exception suggest that this interpretation is correct. 
That the strength of the diaphragm is equal to that of the shell wall is extrapolated from his 
de-mineralised sections which show the continuous nature of these structures. 

Using the joint reports of Leidy and Stump, a composite description of a species emerges: it 
sometimes has a constriction of the neck that separates the anterior third of the shell from the main 
body; at this point an internal diaphragm is present which is pierced by a single, central, circular 
opening. 

Medioli & Scott (1983) do not illustrate the internal division in their specimens but describe it as 
a 'large orifice'. They presume that their material is conspecific with that of Leidy and Stump, 
possibly because all the specimens were collected in America. Fortunately, a single specimen with 
an internal division was found in material kindly left at the British Museum (Natural History) by 
F. S. Medioli and helps to confirm their presumption. The specimen had the following measure- 



GENERA PONTIGULASIA, LA GENODIFFLUGIA AND ZIVKOVICIA 15 

ments: 174 um long, 108 um broad, diameter of aperture 37 urn and diameter of internal opening 
27 um. Furthermore the organic cement pattern of this specimen was typical of that illustrated 
earlier (Ogden, 1983). 

DIAGNOSIS. Shell pyriform, often with a constriction of the neck, most frequently circular in 
cross-section but sometimes slightly compressed, composed mainly of agglutinate mineral 
particles bound by an organic cement; aperture terminal, circular; internally the shell is partitioned 
into two regions by a diaphragm constructed as part of the shell wall but having a single central 
orifice. Type species L. vas (Leidy, 1 874). Three other species are here attributed to the genus on the 
basis of having a single opening in a well constructed diaphragm: L. bryophila (Penard, 1902) (see 
Ogden, 1983 for recent description of this species); L. montana (Ogden & Zivkovic, 1983) and L. 
epiouxi (Chardez & Caspar, 1984). 

Zivkovicia Ogden, 1987 gen. nov. 

This genus was erected by Ogden (1983) to accommodate those species of Pontigulasia which had a 
diaphragm with either one or two internal openings. At that time no distinction was drawn 
between the number of openings in the diaphragm. With the creation of Lagenodifflugia to 
represent those specimens with a single opening, Zivkovicia is redefined here to represent those 
species with two openings. 

In the earlier report (Ogden, 1983), due to an oversight, a type species for Zivkovicia was not 
designated, so under Article 1 3(b) of the International Code of Zoological Nomenclature the genus 
is not taxonomically valid. Nevertheless, the name is still available and to avoid confusion is used 
again here, with an amended diagnosis. 

Although Carter's (1 864) original description ofDiffugia compressa did not include a mention of 
an internal diaphragm, his drawings are so precise that they show the typical V-shaped notch on 
the shell which represents the internal diaphragm. This structure is clearly illustrated by Figs 1 8-25 
of Ogden (1983). It is therefore proposed as the type-species of the genus. The earlier discussion 
(p. 14) relating to the incorrect synonymy of P. vas and P. spectabilis, allows spectabilis to be 
available to include the original description of specimens with a bisected diaphragm (Penard, 1902) 
and the recent description by Ogden (1983), whose P. vas now becomes a synonym of Z. spectabilis. 

DIAGNOSIS. Shell pyriform, often with a distinct constriction of the neck region, either circular or 
compressed in transverse-section, composed mainly of agglutinate mineral particles bound by an 
organic cement matrix; aperture terminal, usually circular; internally the shell is partitioned into 
two parts by an extension of the shell wall to form a diaphragm which is bisected by two circular 
openings. Type species Z. compressa (Carter, 1864), other species Z. spectabilis (Penard, 1902) and 
Z.flexa (Cash & Hopkinson, 1 909). A recent description of the latter species can be found in Ogden 
(1983). 



Discussion 

Associated problems of clearly identifying the internal openings were discussed previously (Ogden, 
1 983). Suffice to say here that both these openings and the aperture can be sealed by an organic cyst 
membrane, and the incidence of a single specimen with a trisected diaphragm in Z. compressa (Fig. 
28, Ogden, 1983) is considered to be an isolated deformity. 

In discussing a possible relationship between Pontigulasia Rhumbler, 1896 and Lesquereusia 
Schlumberger, 1845, Medioli & Scott (1983) suggest that both genera are characterised by a 
constriction at the base of the neck which corresponds to an internal diaphragm, and further state 
that a morphological intergradation exists between the two genera. They consider that the 
remaining difference between the genera, that Lesquereusia is constructed of siliceous idiosomes 
whereas Pontigulasia is always reported to be composed of xenosomes, is insufficient to separate 
these two genera. 

It has already been established by Stump (1936, 1943) that P. vas would not construct a shell or 
reproduce in the absence of extraneous material, even in the presence of abundant food, and that 



16 C. G. OGDEN 

such deprived animals commenced normal reproductive activities when shell making material was 
reintroduced. He concluded that P. vas was unable to secrete its own shell material and suggested a 
possible alternative that individuals might be produced without a shell covering. No reports of 
such naked individuals have been recorded in the literature, but it is well known that some 
agglutinate species are capable of constructing an organic shell, identical to that which in the field 
incorporates mineral particles (Hedley et al., 1976; Netzel, 1972, 1976). Furthermore, the 
deposition of siliceous structures by Lesquereusia spiralis are carried out in the cytoplasm of the 
animal (Harrison et al., 1981), and this species is capable of constructing a shell in the absence of 
extraneous material. It is equally capable of incorporating xenosomes and Stump used this ability 
for creating 'windows' to observe cytoplasmic activity. A new family the Lesquereusiidae was 
designated (Ogden, 1979) to include those members of the Lobosia which secrete their own 
siliceous elements, e.g. Lesquereusia, Netzelia Ogden, 1979, and Quadrulella Cockerell, 1909. 

The suggestion by Medioli & Scott (1983) of considering a relationship between Lesquereusia 
and Pontigulasia is perhaps best treated as an indiscretion on the part of geologists venturing into 
the alien field of biology. Phylogenetic interpretations should be based on the animal as a complete 
organism, which in protozoa would include information on cytoplasmic detail, movement, repro- 
duction as well as external coverings, especially when such information exists in publication and 
the animals are easily collected from the field. 



References 

Carter, H. J. 1864. On freshwater Rhizopoda of England and India; with illustrations. Annals and Magazine 

of Natural History. London. 13(3): 18-39. 
Cash, J. & Hopkinson, J. 1909. The British Freshwater Rhizopoda and Heliozoa. Vol. II. Rhizopoda, part 2. 

The Ray Society, London. 166 pp. 
Cash, J., Wailes, G. H. & Hopkinson, J. 1919. The British Freshwater Rhizopoda and Heliozoa. Vol. IV. 

Supplement to the Rhizopoda. The Ray Society, London. 130 pp. 
Chardez, D. 1985. Note sur les genres Pontigulasia Rhumbler et Zivkovicia Ogden (Rhizopoda, testacea). 

Revue Vervietoise d'Histoire Naturelle 42: 13-16. 
& Caspar, Ch. 1984. Nouveaux thecamoebiens aquatiques du domain des Epioux (Ardenne, Belgique). 

Biologische Jaarb 52: 57-63. 
Harrison, F. W., Dunkelberger, D., Watabe, N. & Stump, A. B. 1981. Ultrastructure and deposition of silica in 

rhizopod amebae. In: T. L. Simpson & B. E. Volcani eds, Silicon and siliceous structures in biological 

systems. Springer- Verlag, New York. pp. 281-294. 
Hedley, R. H., Ogden, C. G. & Mordan, N. J. 1976. Manganese in the shell of Centropyxis (Rhizopodea: 

Protozoa). Cell and Tissue Research 171: 543-549. 
Leidy, J. 1874. Notice of some new freshwater Rhizopods. Proceeding of the Academy of Natural Sciences of 

Philadelphia, ser. 3: 77-79. 
1 879. Freshwater Rhizopods of North America. In: 'United Stated Geological Survey of the Territories', 

Vol. 12. Washington, 324 pp. 
Loeblich, A. R. & Tappan, H. 1964. Thecamoebians'. In: 'Treatise on Invertebrate Paleontology. Part C. 

Protista 2, Vol. 1. C16-C54. The Geological Society of America. 
Medioli, F. S. & Scott, D. B. 1983. Holocene Arcellacea (Thecamoebians) from eastern Canada. Cushman 

Foundation for Foraminiferal Research. Special publication No. 21, 63 pp. 
Netzel, H. 1972. Die Bildung der Gehausewand bei der Thekamobe Centropyxis discoide (Rhizopoda, 

Testacea). Zeitschr if t fur Zellforschung und Mickroskopische Anatomic 135: 45-54. 
1976. Die Abscheidung der Gehausewand bei Centropyxis discoides (Rhizopoda, Testacea). Archivfur 

Protistenkunde 118: 53-91. 
Ogden, C. G. 1979. Siliceous structures secreted by members of the subclass Lobosia (Rhizopoea: Protozoa). 

Bulletin of the British Museum (Natural History) (Zoology) 36: 203-207. 
1983. The significance of the inner dividing wall in Pontigulasia Rhumbler and Zivkovicia gen. nov. 

(Protozoa: Rhizopoda). Protistologica 19: 215-229. 

Penard, E. 1902. Faune Rhizopodique du Bassin du Leman. Geneva. 700 pp. 
Rhumbler, L. 1896. Beitrage zur Kenntnis der Rhizopoden. Zeitschrift fur Wissenschaftliche Zoologie 61: 

38-110. 



GENERA PONTIGULASIA, LAGENODIFFLUGIA AND ZIVKOVICIA 17 

Stump, A. B. 1935. Observations on the feeding of Difflugia, Pontigulasia and Lesquereusia. The Biological 

Bulletin oj the Marine Laboratory , Woods Hole 69: 136-142. 
1936. The influence of test materials on reproduction in Pontigulasia vas (Leidy) Schouteden. The 

Biological Bulletin of the Marine Laboratory, Woods Hole 70: 142-147. 
1943. Mitosis and cell division in Pontigulasia vas (Leidy) Schouteden. Journal of the Elisha Mitchell 

Scientific Society 59: 14-22. 

Manuscript accepted for publication 7 July 1986 

Since this manuscript was submitted for publication the author has become aware of a paper by 
Medioli & Scott (1985), in which they have designated type specimens for certain species. In dealing 
with species of Pontigulasia, Lagenodifflugia and Zivkovicia they make assumptions based solely 
on published descriptions and micrographs of fossil specimens, which do not have information 
relating to the main diagnostic feature the internal dividing diaphragm. In their original paper 
(Medioli & Scott, 1983), they comment on never having been able to satisfactorily observe the 
diaphragm of P. compressa (Carter, 1864), and their failure is again reiterated in Medioli & Scott 
(1985), As neither their figures nor plates illustrate an internal structure it suggests that they 
have failed by both optical and electron microscopy to see this feature in any of their specimens. 
They presume the presence of a diaphragm solely on it corresponding in position to the external 
constriction. Although they admit that this external constriction is often obscured by agglutinate 
material. 

Furthermore, the specimen they selected as neotype of L. vas, from their Maritime Canada 
sample does not even resemble the original selected figure from Leidy (1879). In fact they chose an 
example which was intermediate between Leidy's figured specimen, a smoothly agglutinate form, 
and their extreme cases of coarse agglutination. 

Both of Medioli & Scott's (1983, 1985) papers concerning these genera must be of minimal value 
because of their failure to describe or identify the internal structures which typify these genera. Its 
absence suggests that they have been examining species of Difflugia. Their insistance that the 
diaphragm is absent in fossil forms (p. 35, 1983; p. 29, 1985) of Pontigulasia compressa (Carter, 
1 864) would indicate that they are dealing with species of Difflugia. The construction of the 
diaphragm in this species (Ogden, 1 983) is so robust, being continuous and as thick as the shell wall, 
that even in fractured shells it is the wall that breaks and not the diaphragm. Although often by 
convenience such breaks occur at the junction of the wall and diaphragm. Notwithstanding this, 
specimens clearly identified as Zivkovicia compressa (Carter, 1864) have been recovered from core 
samples taken in Lake Ullswater which had complete diaphragms (Ogden & Ellison, in prep.), and 
in addition possessed the typical organic cement units, described by Ogden (1983), specific to this 
genus. 

Medioli, F. S. & Scott, D. B. 1985. Designation of types, for one genus and nine species of Arcellaceans 
(Thecamoebians), with additional original reference material for four other species. Journal of 
Foraminiferal Research 15: 24-37. 



A revision of the foraminiferal genus Adercotryma 
Loeblich & Tappan, with a description of A. wrightisp. 
nov. from British waters 

P. Bronnimann 

9G, Chemin de Bedex, 1226 Thonex, Geneva, Switzerland 

J. E. Whittaker 

Department of Palaeontology, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 

Adercotryma Loeblich & Tappan (1952) was erected to accommodate Lituola glomerata Brady 
(1878), a species assigned subsequently to Haplophragmium, Haplophragmoides and Trochammina 
by various authors. A recent examination of the type material in the British Museum (Natural 
History), and of specimens from other collections deposited there and in the National Museum of 
Ireland, led to the discovery that the original definition was inadequate and to the recognition of a 
second species. The purpose of this paper is to emend the diagnosis of Adercotryma, to redescribe 
A. glomeratum (Brady), and to describe the new species. 

The generic diagnosis below is based on the redescription of the type species, and differs from the 
original definition in that it recognises the significance of the asymmetrically placed aperture and 
shows the coiling to be trochospiral. Adercotryma is therefore transferred from the Lituolacea to 
the Trochamminacea and placed in a new subfamily of the Trochamminidae. The definition 
follows the format adopted by Bronnimann et al. (1983) in their reclassification of the 
Trochamminacea. 

Superfamily TROCHAMMINACEA Schwager, 1877 

Family TROCHAMMINIDAE Schwager, 1877 

Subfamily ADERCOTRYMINAE nov. 

DEFINITION. Test free; adult, a completely or almost completely involute, cone-like, trochospire; 
wall agglutinated, imperforate, single-layered, aperture interiomarginal, single; without secondary 
septa or infoldings of the umbilical chamber walls or incomplete secondary partitions. 

TYPE GENUS. Adercotryma Loeblich & Tappan, 1952. 

REMARKS. The Adercotryminae differs from all other subfamilies in that its members are 
completely or almost completely involute on the spiral side. Bronnimann et al. (1983: 204) 
distinguished the Trochamminacea from the Ataxophragmiacea on the ratio of spire height to 
umbilical diameter: the former being always smaller than the latter in the Trochamminacea. At 
first sight, the high cone-shaped test of Adercotryma does not fulfil this criterion, but since the 
coiling is involute and the proloculus is situated within the shell (see Figs 3, 6), the spire height 
measured from the proloculus is invariably less than the umbilical diameter. 

Genus ADERCOTRYMA Loeblich & Tappan, 1952 

TYPE SPECIES. Lituola glomerata Brady, 1878. Recent, marine; distribution apparently worldwide. 
Lectotype from Arctic waters. 

EMENDED DEFINITION. Test free; coiling trochospiral, adult an inverted cone, completely or almost 
completely involute on both sides. Chambers axially elongate. Aperture single, interiomarginal, 
umbilical, symmetrical with respect to long axis of chamber. Wall agglutinated, single layered, 
imperforate. 



Bull. Br. Mm. nat. Hist. (Zool.) 52(1): 19-28 Issued 29 January 1987 



20 



P. BRONNIMANN & J. E. WHITTAKER 



REMARKS. The slit-like aperture rests with its border on the first and on the penultimate chamber of 
the final whorl (Paratrochammina-type aperture). Adercotryma differs from Paratrochammina 
Bronnimann, 1979 (type species: P. madierae Bronnimann, 1979) and all other genera of the 
Trochammininae by its spirally involute enrolment, axially elongate chambers, symmetrical 
interiomarginal aperture (with respect to the long axis of the chamber), and inverted cone-like test. 

Adercotryma glomeratum (Brady) 
Figs 1, 2A-F, 3 A, 4A-E, 5A-J, 6A-F 

1878 Lituola glomerata Brady: 433, pi. 20, figs la-c. 

1 884 Haplophragmiwn glomeratum (Brady); Brady: 309, pi. 34, figs 15-18. 

1910 Haplophragmoides glomeratum (Brady) (sic); Cushman: 104, figs 158-161 (after Brady, 1884). 

1 93 1 Trochammina glomerata (Brady); Wiesner: 1 1 2, pi. 1 7, figs 204, 205. 

1952 Adercotryma glomerata (Brady) (sic); Loeblich & Tappan: 141, figs 1-4. 

1961 Adercotryma glomerata glomerata (Brady); Saidova: 35, pi. 10, fig. 54. 

1961 Adercotryma glomerata abyssorum Saidova (sic): 36, pi. 10, fig. 55. 

1975 Adercotryma glomerata antarctica Saidova (sic): 75, pi. 96, fig. 6. 

MATERIAL. Extant material in the Brady Collection of the British North Polar Expedition 
(1875-1876), labelled Lituola glomerata, is as follows: Station A, off Tyndall Glacier, 27fms 
(49m); F, between Walrus Shoal and Victoria Head, 57fms (104m); H, Franklin Pierce Bay, 
13-1 5 fms (24-28 m); I, Allman Bay, 25 fms (46m); J, Dobbin Bay, 45^17 fms (82-86 m);. K, 
Dobbin Bay, 1 1 3 fms (207 m); N, off Hayes Point, 35 fms (64 m); O, off Cape Frazer, 50 fms (92 m) 
and P, off Cape Frazer, 80 fms (146 m). These localities are from the northern part of Baffin Bay 
and Smith Sound (between Ellesmere Island, NE Canada, and W Greenland). All the slides 
contain a few specimens at least, and some (e.g. station G) as many as 50. 




Fig. 1. Adercotryma glomeratum (Brady). Paralectotype, 1955.10.28.1732. Interpretative drawing of 
specimen in Figs 6A-C, taken at the third level of dissection (see explanation of Fig. 6C), showing 
chambers 1 to 9. Hatched areas represent exposed walls of earliest chambers, x 300. 

From Franklin Pierce Bay, lat. 7928'N, station H, depth 46 fathoms (84 m). British North Polar 
Expedition of 1875-1876, ex BMNH slide no. 1955.10.28.1731-1780, labelled 'Lituola glomerata 
Brady'. 



GENUS ADERCOTRYMA 



21 









Fig. 2A-F. Adercotryma glomeratum (Brady). 2A-C, Paralectotype, 1955.10.28.1701. Interpretative 
drawing of specimen in Figs 5E-G, J. 2A, umbilical view showing the overlapping chamber walls and 
the preserved apertural slits of the last three chambers; 2B, spiral view; 2C, edge view showing aperture 
of final chamber, with axis of coiling marked by line A. 2D-F, Paralectotype, 1955.10.28.1700. 
Interpretative drawing of specimen in Figs 5A-D. 2D, umbilical view showing aperture of final 
chamber in part masked by agglutinated or secreted material; 2E, spiral view; 2F, edge view of inverted 
cone-like test with aperture of final chamber in part masked, axis of coiling is indicated by line A. Both 
x!50. 

Both from slide labelled ' Lituola glomerata Brady'. British North Polar Expedition of 1875-1876. 
Cape Frazer, lat. 7945'N, station O, depth 50 fathoms (92m), ex BMNH slide no. 
1955.10.28.1700-1731. 



LECTOTYPE. 1955.10.28.1781 (Figs 4A-E). From Brady's syntypic series, obtained from Station P, 
off Cape Frazer, Arctic Canada, depth 80 fathoms (146 m). Believed to be the specimen figured by 
Brady (1878, pi. 20, fig. Ib). 

DESCRIPTION (LECTOTYPE). Test free; a dextral, tightly coiled trochospire, with 4 chambers in the 
final whorl, each gradually increasing in size; involute on spiral side. Test a short, broad, inverted 
cone-like structure, flatly truncated spirally, rounded-convex umbilically, broadly rounded peri- 
pherally and somewhat rounded laterally. In edge view, 3 chambers seen on both sides. In spiral/ 
umbilical view, oval-lobate; umbilical side with a small, well-defined subcircular and shallow 
axial depression. Adult chambers much elongated in axial (edge) view, narrow radially and some- 
what elongate tangentially, more inflated towards the spiral, than towards the umbilical side. 
Intercameral sutures straight but indistinct spirally; straight, distinct, laterally and umbilically. 
Aperture single, interiomarginal, a narrow elongate slit with rounded extremities, at umbilical end 
of chamber; symmetrical with respect to its long axis. Border of aperture rests on the first and 



22 



P. BRONNIMANN & J. E. WHITTAKER 





Fig. 3A Adercotryma glomeratum (Brady). Paralectotype, 1955. 10.28. 1782. Section cut parallel to axis 
of coiling. Note the thin-walled proloculus already slightly elongate in the direction of the coiling axis. 
x205. 

From Cape Frazer, lat. 7945'N, station P, depth 80 fathoms (146m). British North Polar 
Expedition of 1875-1876, ex BMNH slide no. 1955.10.28.1781-1799, labelled 'Lituola glomerata 
Brady'. 

Fig. 3B. Adercotryma wrighti Bronnimann & Whittaker sp.nov. ZF 4453. Section cut slightly obliquely 
to axis of coiling. x250. 

From south of Mull, W Scotland, depth 20 fathoms (37m). S.Y. Runa station 2, collected 1913. 
Heron-Allen & Earland Collection (BMNH), slide labelled ' Haplophragmium glomeratum (Brady)'. 



penultimate chambers of final whorl (Paratrochammina-type). Final chamber covers about half of 
the preceding apertural slit. Wall agglutinated, imperforate, coarser on truncated, spiral side than 
on rounded-comvex, umbilical side. Colour, prior to coating for SEM photography, yellowish- 
brown. 

DIMENSIONS (LECTOTYPE). Maximum spiral/umbilical diameter 320 um, minimum diameter 
270 um, height 260 um. Height of apertural slit c. 12 um. 

PARALECTOTYPES: 3 sinistral specimens (1955.10.28.1700-1702) are figured in Figs 2A-F, 5A-J; 
another 3 (1955.10.28.1732, 1955.10.28.1783 and 1955.10.28.1703), dissected out to show various 
aspects of the internal coiling, are figured in Figs 1, 6A-F, whilst a further paralectotype 
(1955.10.28.1782) has been sectioned and is illustrated in Fig. 3 A. These specimens, as with others 
remaining in Brady's syntypic series, vary considerably in their dimensions, elongation of the 
chambers in the final whorl, depression of the sutures, depth of the umbilicus and spiral aspect. For 
further comments, see the figure explanations and Remarks section below. The maximum spiral/ 
umbilical diameter of the figured paralectotypes varies from 230 to 290 um, the test height, from 
250 to 270 um. 

REMARKS. Brady's small and enigmatic species was placed by authors in Lituola, Haplophragmium, 
Haplophragmoides and Trochammina prior to the erection of Adercotryma by Loeblich & Tappan 
( 1 952). The generic changes stem mainly from differing interpretations of the mode of coiling of the 
curious cone-shaped test. 

Although Brady (1878) originally referred to the test as merely '. . . spiral in arrangement', his 
subsequent comparison (Brady, 1884) of the overall shape with that of a'. . . nautiloid species, such 
as Haplophragmium latidorsatum, drawn out as the umbilici so as to form a test bearing some 
resemblance to the oval Alveolinae', implied that the coiling was planispiral. He clearly was not 
sure, however, as he made much in these two papers of the unusual 'unsymmetrical convolutions'. 

It was Cushman (1910) who first described the coiling, without reservation, as planispiral, 
placing Brady's species in his new genus Haplophragmoides, an assignment which was generally 



GENUS ADERCOTRYMA 




Fig. 4A-E. Adercotryma glotneratum (Brady). Lectotype, 1955.10.28.1781. 4A-D, spiral, edge 
(apertural), umbilical and edge (antapertural) views, x 1 50. 4E, detail of lateral, open part of aperture, 
x525. 

From slide labelled ' Lituola glomeratum Brady'. British North Polar Expedition of 1 875-1 876. Cape 
Frazer, lat. 7945'N, station P, depth 80 fathoms (146 m), ex BMNH slide no. 1955.10.28.1781-1799. 



followed for over forty years. The only exception was Wiesner (1931) who placed glomerata in 
Trochammina, although he did not make a detailed examination of its morphology and his paper 
offers no evidence for trochospiral coiling. A year earlier, however, Lacroix (1930) had considered 
the position of the aperture, ignored completely by Cushman, to be more in keeping with a 
trochospiral genus. In terms of coiling, Lacroix considered Brady's species transitional between 
the planispiral Haplophragmoides and the trochospiral Trochammina, but nevertheless retained it 
in the former genus. 

In 1952, Loeblich & Tappan erected a new lituolid genus Adercotryma, with Lituola glomerata 
Brady as type. The name refers to the apertural features, derived from two Greek words aderco- 
unseen, invisible, and tryma- meaning a hole or aperture. The gender of the name Adercotryma is 
neuter, and the specific name should be construed as glomeratum, not glomerata as originally 
written. Loeblich & Tappan (1952) distinguished their new genus from Haplophragmoides on the 
somewhat asymmetrical, completely involute, rather than slightly evolute test which has its 
greatest dimension in the axis of coiling, and by its aperture which lies near the umbilicus of 
one side, rather than in the plane of coiling at the periphery. Of these features, only two are 
fundamentally different from those of Haplophragmoides: the asymmetrical test morphology and 
the asymmetrical interiomarginal apertural position. In no part of their original paper, nor in 1964, 
did Loeblich & Tappan discuss the curious asymmetry of what they obviously assumed to be a 
planispiral test. Adercotryma was placed in the Haplophragmoidinae Maync, 1952 (Lituolidae de 
Blainville, 1825), in which were included both planispiral and streptospiral forms. 

Even though the test of A. glomeratum is involute, the external and internal morphology clearly 



24 



P. BRONNIMANN & J. E. WHITTAKER 




Figs 5A-J. Adercotryma glomeratum (Brady). 5A-D, Paralectotype, 1955.10.28.1700. Spiral edge, 
oblique-umbilical and umbilical views, x 150. 5E-G, J, Paralectotype, 1955.10.28.1701, E-G, spiral, 
edge and umbilical views, x 1 50; 5J, detail of interiomarginal apertures of final and penultimate 
chambers, in umbilical view, x475. 5H, I, Paralectotype, 1955.10.28.1702. Spiral and edge views, 
x!50. 

All specimens from slide labelled 'Lituola glomerata Brady'. British North Polar Expedition of 
1875-1876. Cape Frazer, lat. 7945'N, station O, depth 50 fathoms (92m), ex BMNH slide no. 
1955.10.28.1700-1731. 



Figs 6A-F. Adercotryma glomeratum (Brady). 6A-C, Paralectotype, 1955. 10.28. 1 732. Stereo-pairs of 
three stages of dissection, perpendicular to axis of coiling; the involute spiral side has been removed. 
The third stage of dissection (6C) has broken open the earliest whorl and proloculus (see Fig. 1, for 
interpretative drawing). 6D, E, Paralectotype, 1955.10.28.1783. Stereo-pairs of dissected specimen 
shown at two different tilts. Dissection is in plane virtually parallel to coiling axis. 6F, Paralectotype, 
1955.10.28.1703. Stereo-pair of specimen dissected perpendicular to axis of coiling. All x 150. 

Figs 6A-C from Franklin Pierce Bay, lat. 7928'N, station H, depth 46 fathoms (84 m), ex BMNH 
slide no. 1955.10.28.1731-1780. Figs 6D, E from Cape Frazer, lat. 7945'N, station P, depth 80 
fathoms (146m), ex BMNH slide no. 1955.10.28.1781-1799. Fig. 5F, same locality, station O, depth 
50 fathoms (92m), ex BMNH slide no. 1955.10.28.1700-1731. British North Polar Expedition of 
1875-1876. 



GENUS ADERCOTRYMA 



25 




26 



P. BRONNIMANN & J. E. WHITTAKER 




Figs. 7A-J. Adercotryma wrighti Bronnimann & Whittaker sp.nov. 7A-D, Holotype, NMI no. 
149. 1985. Spiral, edge (apertural), umbilical and edge (antapertural) views. 7E, F, Paratype, NMI no. 
4.1980. Oblique-umbilical and edge views. 7G-J, Paratype, NMI no. 5.1980. Spiral, edge (apertural), 
edge (antapertural) and umbilical views. All x 175. 

All from offDrogheda, E Ireland, depth 16 fathoms (29m); ex slide no. 34, labelled 'Dublin: off 
Drogheda, 16 fms & Lambay Deep, 70 fms (mixed)', J. Wright Collection, 13-1921, National Museum 
of Ireland. 



indicates a trochospiral mode of coiling (Figs 1-6). As well as the asymmetrical aperture, the adult 
test, when orientated with the axis of coiling in vertical position, shows a truncated, more or less 
flattened aboral or spiral side, and an obtusely pointed, ovoid-rounded oral or umbilical side. This 
differentiation is typical of trochospiral tests. 

Saidova (1961 ; 1975) introduced two new subspecies of A. glomeratum, namely A. g. abyssorum 
and A. g. antarctica, respectively. They are figured together with a typical A. g. glomeratum also in 
Saidova ( 1 975, pi. 96, figs 4-6). The difference in shell morphology said to characterise the two (test 
size, chamber shape and elongation) falls within the range of variation seen in our paralectotypes 
(compare our Figs 5B and 51 with Saidova's pi. 96, figs 6 and 5, respectively), whilst the type of 



GENUS ADERCOTRYMA 27 

agglutinant, also used by Saidova (1961) as a distinguishing feature of A. g. abyssorum, is not 
considered by us to have any taxonomic validity. 

A.glomeratum (Brady) differs from A . wrighti sp. nov. in having a broadly inverted cone-like test 
with 4 axially elongate chambers in the final whorl, in the shape of the adult chambers, and in the 
apertural features. See also pp 27, 28 for further remarks on their differences. In the material 
studied, A. glomeratum always has 4 chambers in the final whorl, even the preceding whorl (Figs 1, 
6C) has 4 chambers. This results in 3 chambers being visible on either side of the test when seen in 
edge view (aperturally and antaperturally). 

A. glomeratum is a very wide-ranging species both in terms of latitudinal and depth distribution 
as noted by Saidova (1975) and Culver & Buzas (1985). 

Adercotryma wrighti Bronnimann & Whittaker sp. nov. 
Figs 3B, 7A-J 

1908 Haplophragmium glomeratum (Brady); Millett: 5 (list), pi. 1, fig. 6 (non Lituolaglomerata Brady, 1878). 
1913 Haplophragmium glomeratum (Brady); Heron- Allen & Earland: 46, pi. 2, fig. 14. 

DIAGNOSIS. A species of Adercotryma with only 3 chambers in the final whorl. In spiral/umbilical 
view, test oval-lobate, maximum diameter often almost twice the minimum diameter. In edge view, 
3 chambers seen on apertural side, only 2 on antapertural side. Aperture single, interiomarginal, a 
bilobed narrow, elongate slit without rounded extremities at umbilical end of chamber. 

NAME. In honour of Joseph Wright, in whose collection from Dublin Bay this species was first 
noticed. 

HOLOTYPE. National Museum of Ireland (NMI) no. 149.1985. Illustrated in spiral, edge 
(apertural), umbilical and edge (antapertural) views in Figs 7A-D. Ex J. Wright Collection, slide 
34. 

TYPE LOCALITY. OffDrogheda, E Ireland, depth 16 fathoms (30 m). 

DESCRIPTION (HOLOTYPE). Test free; a dextral, tightly coiled trochospire, with 3 chambers in the 
final whorl, gradually increasing in size; involute on spiral side. Test an inverted cone-like struc- 
ture, truncated spirally, rounded-convex umbilically and broadly rounded peripherally. In edge 
view, 3 chambers seen on apertural side, 2 on antapertural side. In spiral/umbilical view, oval- 
lobate, maximum diameter almost twice the minimum diameter; umbilical side with very shallow 
and small axial depression. Adult chambers elongate in axial (edge) view, less elongate in tangential 
direction, narrow in radial direction; inflated equally both spirally and umbilically. Intercameral 
sutures well defined and slightly incurved laterally and umbilically, less well defined and straight 
spirally. Aperture single, interiomarginal, a bilobed narrow slit without rounded extremities, at 
umbilical end of chamber, symmetrical with respect to long chamber axis. Border of aperture rests 
on first and on penultimate chamber of final whorl (Paratrochammina-typo). Wall agglutinated, 
imperforate, coarser on spiral side than on umbilical side. Colour of test, prior to coating for SEM 
photography, orange-brown. 

DIMENSIONS (HOLOTYPE). Maximum spiral/umbilical diameter 240 um, minimum diameter 1 50 um, 
height 220 um. 

PARATYPES. Two paratypes are figured herein. NMI no. 4. 1980 (Figs 7E, F) is a sinistral specimen; 
the aperture is perfectly preserved and shows the bilobed, narrow, elongate slit; the development of 
a central, triangular lip-like projection of the chamber wall serving to divide the aperture into two 
virtually identical parts. This specimen has a maximum spiral/umbilical diameter of 220 um and 
test height of 240 um. NMI no. 5.1980, the other illustrated paratype (Figs 7G-J), is dextrally 
coiled like the holotype. Its maximum spiral/umbilical diameter is 200 um, the test height 210 um. 
The sectioned specimen (ZF 4453), from the Heron-Allen & Earland Collection, off W Scotland, 
has a maximum diameter of 220 um and height of 180 urn; it is figured in Fig. 3B. 

REMARKS. Adercotryma wrighti sp. nov. is easily distinguished from A. glomeratum (Brady) by the 
overall shape of the test, only 3 chambers in the final whorl, the shape of the adult chambers and the 



28 P. BRONNIMANN & J. E. WHITTAKER 

bilobed apertural features formed by the triangular lip-like projection of the chamber wall. A. 
glomeratum always has 4 chambers in the final whorl and a test which in spiral/umbilical aspect has 
a maximum diameter little greater than the minimum. 

Both Millett (1908) and Heron- Allen & Earland (1913) show specimens from W Ireland with 
only 3 chambers in the final whorl. We have examined their collections and many slides in the 
Brady and the Norman Collections (BMNH), labelled Haplophragmium glomeratum (Brady), 
from Scotland, Ireland and N England and all exclusively contain A. wrighti rather than A. 
glomeratum. Careful study of specimens previously recorded as A. glomeratum may extend the 
present range of A. wrighti beyond the British Isles. 

Acknowledgements 

The Director of the National Museum of Ireland is thanked for his permission to borrow and 
photograph specimens from the Wright Collection; Mr J. M. C. Holmes facilitated the loan. We 
are pleased to acknowledge the technical skill of Mr R. L. Hodgkinson, British Museum (Natural 
History), in preparing the dissections and thin sections of the Adercotryma spp.; Mrs L. M. 
McCormick and Mr P. V. York took the SEM and optical photographs, respectively, whilst Drs 
C. G. Adams and M. K. Howarth, also of the same institution, kindly read the manuscript and 
suggested many improvements. The research of P. Bronnimann is in part funded by the Fonds 
National Suisse. 

References 

Blainville, H. M. D. de 1 825. Manuel de Malacologie et de Conchy Uologie. 664 pp., 87 pis. F. G. Levrault, Paris. 
Brady, H. B. 1878. On the reticularian and radiolarian Rhizopoda (Foraminifera and Polycystina) of the 

North Polar Expedition of 1875-76. Ann. Mag. nat. Hist., London, ser. 5, 1: 425-440, pis 20, 21. 
1884. Report on the Foraminifera dredged by H.M.S. Challenger during the years 1873-76. Rep. sclent. 

Results Voy. Challenger (Zool.), London, 9: 1-814, pis 1-1 15. 
Bronnimann, P. 1979. Recent benthonic foraminifera from Brasil Morphology and Ecology; part 4: 

Trochamminids from the Campos Shelf with description of Paratrochammina n.gen. Palaeont. Z., 

Stuttgart, 53: 5-25, figs 1-10. 
, Zaninetti, L. & Whittaker, J. E. 1983. On the classification of the Trochamminacea (Foraminiferida). J. 

foramin. Res., Washington, 13: 202-218, pis 1-3. 
Culver, S. J. & Buy as, M. A. 1985. Distribution of Recent benthic foraminifera of the North American Pacific 

Coast from Oregon to Alaska. Smithson. Contrib. Mar. Sci., Washington, 26: 1-234, figs 1-139. 
Cushman, J. A. 1910. A monograph of the Foraminifera of the North Pacific Ocean; Part 1 , Astrorhizidae and 

Lituolidae. Bull. U.S. natn. Mus., Washington, 71(1): 1-134, figs 1-203. 
Heron-Allen, E. & Earland, E. 1913. The Foraminifera of the Clare Island District, County Mayo, Ireland. 

(Clare Island Survey, part 64.) Proc. R. Ir. Acad., Dublin, 31: 1-188, pis 1-13. 
Lacroix, E. 1930. Les Lituolides de plateau continental mediterranean entre Saint-Raphael et Monaco. Bull. 

Inst. oceanogr. Monaco 549: 547-550, figs 1-21. 
Loeblich, A. R. Jr & Tappan, H. 1952. Adercotryma, a new Recent foraminiferal genus from the Arctic. /. 

Wash. Acad. Sci., 42: 141-142, figs 1-4. 
& 1964. Protista 2. Sarcodina chiefly 'Thecamoebians' and Foraminiferida. In Moore, R. C. (ed.) 

Treatise on Invertebrate Paleontology, Part C, 1: 1-510, figs 1-399. University of Kansas Press. 
Maync, W. 1952. Critical taxonomic study and nomenclatural revision of the Lituolidae based upon the 

prototype of the family, Lituola nautiloidea Lamarck, 1804. Contrib. Cushman Lab. foramin. Res., Sharon, 

3: 35-56, pis 9-12. 

Millett, F. W. 1908. The Recent Foraminifera ofGalway. 8 pp, 4 pis. W. Brendon & Son, Plymouth. 
Saidova, Kh. M. 1961. Foraminiferal ecology and palaeogeography of the far eastern seas of the USSR and 

northwestern part of the Pacific Ocean. 232 pp, 31 pis. Akad. Nauk SSSR, Inst. Okeanologii, Moskow. (In 

Russian.) 
1975. Benthonic foraminifera of the Pacific Ocean. 3 vols. 875 pp, 116 pis. Akad. Nauk SSSR, Inst. 

Okeanol. P.P. Shirshova, Moskow. (In Russian.) 
Schwager, C. 1877. Quadro del proposto sistema di classificazione dei foraminiferi con guscio. Boll. R. com. 

geol. Ital., Florence, 8: 18-27, pi. 1. 
Wiesner, H. 1931. Die Foraminiferen der Deutschen Siidpolar-Expedition 1901-1903. Dt. Sudpol.-Exped., 

Berlin (Zool.), 20: 49-165, pis 1-24. 

Manuscript accepted for publication 15 January 1986 



Hermit crabs associated with the bryozoan 
Hippoporidra in British waters 

J. D. D. Bishop 

Departments of Palaeontology and Zoology, British Museum (Natural History), Cromwell Road, 
London SW7 5BD 

The cheilostome bryozoan Hippoporidra lusitania is not associated exclusively with the 
hermit crab Pagurus cuanensis as some previous reports have implied. It has been found 
with two other species of pagurid, Anapagurus chiroacanthus and Cestopagurus timidus; its 
occurrence with P. cuanensis requires confirmation. 

Species of the cheilostome bryozoan genus Hippoporidra preferentially or exclusively encrust 
gastropod shells inhabited by hermit crabs (Crustacea: Anomura: Paguroidea). Helicospiral 
growth of an established bryozoan colony may extend the crab's domicile well beyond the aperture 
of the original gastropod shell (Taylor & Cook, 198 1). The type-species of the genus, Hippoporidra 
edax (Busk), was first described as a fossil from the Coralline Crag (Pliocene) of eastern England, 
but the name has subsequently been used for living specimens from both sides of the North 
Atlantic. However, material from Recent British seas that had formerly been referred to H. edax 
was distinguished as a new species, H. lusitania, by Taylor & Cook (1981). 

No details were given by Taylor & Cook (1981) of the hermit crabs with which H. lusitania 
occurs. A few earlier records of Recent Hippoporidra edax from Britain, which may be assumed to 
refer to H. lusitania, mentioned Pagurus cuanensis Bell as the associated pagurid. Thus, Moore 
(1937) reported a single colony of the bryozoan with P. cuanensis collected off the Isle of Man. This 
record was repeated by Bruce et al. (1963) and quoted by Cook (1964). Eggleston (1972) reported 
that in Manx waters Hippoporidra was in fact restricted to P. cuanensis, and this apparent example 
of extreme stenotopy in a bryozoan was quoted by Ryland (1976). Hay ward & Ryland (1979) gave 
P. cuanensis as the preferred species of British Hippoporidra, and did not name any other pagurid 
with which the bryozoan was found. 

The Bryozoa collection of the Zoology Department of the British Museum (Natural History) 
contains 21 colonies of//, lusitania, including the type series from Guernsey and part of the Manx 
material studied by Eggleston. The associated hermit crab is present in only four examples, as 
detailed in Table 1 . 

It is clear from this that H. lusitania is not restricted to Pagurus cuanensis, even off the Isle of 
Man. Indeed, its occurrence with P. cuanensis at all requires confirmation. P. cuanensis reaches a 
considerably larger size than either Anapagurus chiroacanthus (Lilljeborg) or Cestopagurus timidus 



Table 1 


Specimen 






H 


W Hermit crab 


Isle of Man 


D. Eggleston 


1963.12.30.1 


11-0 


11-5 Anapagurus chiroacanthus 


Guernsey 


A. M. Norman 


1911. 10.1. 1143H 


12-0 


10-0 Anapagurus chiroacanthus 






(Paratype) 






Guernsey 


A. M. Norman 


1911.10.1.11431 


8-5 


8-5 Anapagurus sp. (fragment) 






(Paratype) 






Scilly Isles 


M. H. Thurston 


1965.8.18.26 


7-5 


4-5 Cestopagurus timidus 



H = approximate height of gastropod/bryozoan measured along axis, in mm. 
W = approximate 'body whorl' diameter of gastropod/bryozoan, in mm. 



Bull. Br. Mus. not. Hist. (Zool.) 52(1): 29-30 



Issued 29 January 1987 



30 J. D. D. BISHOP 

(Roux), the carapace lengths given by Bouvier (1940) being 8-12 mm, 46 mm and 4-5 mm respec- 
tively. All H. lusitania colonies available for study at the BM(NH) are relatively small; the largest 
(Plymouth, T. Hincks, 1 899.5. 1.1517) has a height of c. 17 mm and a 'body whorl' diameter of c. 
14 mm. It therefore seems probable from the limited material available that Hippoporidra lusitania 
may be most commonly associated with relatively small hermit crabs. 



Acknowledgements 

I wish to thank R. W. Ingle and P. L. Cook for help with the pagurids and bryozoans respectively. 

References 

Bouvier, E.-L. 1940. Decapodes marcheurs. Faune de France 37: 1^04. 

Bruce, J. R., Colman, J. S. & Jones, N. S. 1963. Marine fauna of the Isle of Man and its surrounding seas. 

L.M.B.C. Memoirs on Typical British Marine Plants and Animals 36: i-ix and 1-307. 
Cook, P. L. 1964. Polyzoa from West Africa. Notes on the genera Hippoporina Neviani, Hippoporella Canu, 

Cleidochasma Harmer and Hippoporidra Canu & Bassler (Cheilostomata, Ascophora). Bulletin of the 

British Museum (Natural History), Zoology series 12: 1-35. 
Eggleston, D. 1972. Factors influencing the distribution of sub-littoral ectoprocts off the south of the Isle of 

Man (Irish Sea). Journal of Natural History 6: 247-260. 
Hay ward, P. J. & Ryland, J. S. 1979. British ascophoran bryozoans. Synopses of the British Fauna (New 

Series) 14: i-v and 1-312. 
Moore, H. B. 1 937. Marine Fauna of the Isle of Man. Proceedings and Transactions of the Liverpool Biological 

Society 50: 1-293. 

Ryland, J. S. 1976. Physiology and ecology of marine bryozoans. Advances in Marine Biology 14: 285-443. 
Taylor, P. D. & Cook, P. L. 1981. Hippoporidra edax (Busk 1859) and a revision of some fossil and living 

Hippoporidra (Bryozoa). Bulletin of the British Museum (Natural History) , Geology series 35: 243-25 1 . 

Manuscript submitted for publication 6 September 1985 



The first zoea of three Pachygrapsus species and of 
Cataleptodius floridanm (Gibbes) from Bermuda and 
Mediterranean (Crustacea: Decapoda: Brachyura) 

R. W. Ingle 

Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD 

Introduction 

Affinities of larvae belonging to the family Grapsidae have been reviewed by Aikawa (1929), Wear 
(1970), Rice (1980) and Wilson (1980). Within the four subfamilies composing this family 'the 
larval development of less than 15% of all the species has been described' (Wilson, 1980: 756). 
Many descriptions are insufficient for meaningful comparative studies and, because of the 
apparent difficulty of rearing these small zoeae, a large percentage of studies describe only the first 
stage. 

Wilson (1980) has provided a useful and comprehensive table of seven comparative features of 
the first stage zoeae of 47 grapsid species. To this list may be added the following accounts which 
contain more or less adequate details for comparative purposes. Plagusiinae: IPlagusia depressa, 
Rice & Williamson, 1977. Varuninae: Eriocheir japonica, Gaetice depressus, Hemigrapsus longi- 
tarsus, H . penicellatus , H. sanguiensis , Terada, 1981. Grapsinae: Metopograpsus latifrons, Kakati, 
1982; M. messor, Rajabai, 1962. Sesarminae: Aratus pisonii, Hartnoll, 1965; Chasmagnathus 
convexus, Saba, 1974; C. laevis, Helograpsushaswellianus, Green & Anderson, 1973; Metasesarma 
rousseauxi, Rajabai, 1962; Sesarma erythrodactyla, Green & Anderson, 1973; S. perracae, Soh 
Chen Lam, 1969; S. tetragonum, Rajabai, 1962. 

Within the genus Pachygrapsus larval stages are known for only three of the fifteen or so 
accepted species (viz. P. marmoratus, P. transversus, P. crassipes). Of these, the complete develop- 
ment has been described for P. marmoratus (Fabricius) and P. crassipes Randall. Larval descrip- 
tions of P. marmoratus are based, except for the first stage, upon plankton collected material (see 
Cano, 1892; Hyman, 1924; Bourdillon-Casanova, 1960), but some of Cano's figures, also repro- 
duced by Hyman, may not even be of a Pachygrapsus (see p. 000). Laboratory hatched first stage 
zoea of P. transversus was described superficially by Lebour (1944) and Rossignol (1957) identified 
a plankton caught megalopa to this species. Villalobos (1971) described the first zoeal stage of 
P. crassipes. This species was laboratory reared to fifth zoeal stage by Schlotterbeck (1976) and a 
plankton caught megalopa was tentatively assigned to P. crassipes by Rathbun (1923). 

During 1973 first stage zoeae of Pachygrapsus marmoratus were obtained from a laboratory held 
crab collected by R. B. Manning off the coast of Tunisia and in 1983 the first stage zoeae of P. 
gracilis and of P. transversus were hatched from crabs held by the author in the Biological Station, 
Bermuda. Although the larvae of these species were not reared beyond the first zoeal stage it would 
seem desirable to give an account of this material to supplement meagre larval information at 
present available on this genus and also to compare (see Table) the first stages of these four 
Pachygrapsus species. Opportunity is also taken to describe the first stage zoea of the xanthid 
Cataleptodius floridanus (also hatched at Bermuda), the larvae of which were studied by Kurata 
(1970) but whose account was never published (see Martin 1984: 233, footnote). 



Materials and methods 

The first zoea of P. marmoratus was hatched from a crab collected in the canal leading from 



Bull. Br. Mus. nat. Hist. (Zool.) 52(1), 31^*1 Issued 29 January 1987 



R. W. INGLE 

























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P. crassipes 
(Schlotterbec] 


0-90 mm 


denticles 
conspicuous 


en 


two small acu 


denticles only 
distal part 


oo 


slightly latera 
expanded, wi1 


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alobos(1971); 2 Leboi 
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Jourdillon-Casanova 
:se as very conspicuo 




P. tranversus 
(present material) 


~E 

E 

o 

o 


denticles 
inconspicuous 




denticles numerous 


long and subacute 
on distal part 


i 


laterally expanded, 
with obtuse 8 dorso- 


lateral processes 


minute, one larger 
than 10 the other 


iffers from the one given by VilL 
ur shows 3 setae; *Bourdillon- 
ses shown on this segment by I 
tourdillon-Casanova shows the 


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denticles 
conspicuous 




denticles few and 


obtuse on distal 
part 


ON 


laterally expanded 
with obtuse dorso- 


lateral processes 


both very minute 


tures by which his ace 
cles as conspicuous; 
te; 9 the conspicuous 
hese equally develop 


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Table 1 . Comparati 


FEATURE 


j' 
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Carapace, posterio- 
margins 


Antennule, aesthete 


Antenna, spinous p 


investment 


Maxilla, coxal endi 


Abdomen, fourth s< 




Telson, lateral spini 


'Schlotterbeck tabula 
H960) gives 1-10 mm 
^Lebour shows these 
margins (see Fig. 3h i 
specimens examined. 



FIRST ZOEA 33 

southern Punic Port, Salammbo, Tunisia, 18.7.1973. Ovigerous Pachygrapsus gracilis and Cata- 
leptodiusfloridanus were collected at various localities from the intertidal mud flats at Ferry Reach 
and P. transversus from beneath stones at Whalebone Bay, Bermuda, all in September/October 
1983. The eggs hatched within 2-3 days of the crab's confinement in aerated aquaria water held at 
20-24C and the larvae were fed newly hatched Anemia nauplii. Live zoeae were subsequently 
transported to a rearing laboratory at the British Museum (Natural History) in London but none 
survived to the second stage. 

Measurements given are: T.T. = distance between tips of dorsal and rostral spines; C.L.= 
carapace length from between the eyes to the posterio-lateral margin. The material has been 
incorporated in the Collections of the British Museum (Natural History), accession number: 1985: 
463^68. 



Descriptions 

Family GRAPSIDAE MacLeay, 1838 
Subfamily GRAPSINAE MacLeay, 1838 

Pachygrapsus gracilis (de Saussure, 1858) 

Dimensions: T.T. 0-60-0-70 mm. C.L. 0-35-0-38 mm. 

Carapace (Fig. la): dorsal spine short and straight, stout proximally, slightly more than one third carapace 

length; rostral spine of moderate length and stout; dorso-median elevation prominent; at least four pairs of 

anterio-median setules and a pair of posterio-median setules present; posterio-lateral margins of carapace 

narrowly rounded with 3-4 obtuse denticles and lateral microscopic setules (inset to fig.). 

Eyes: partly fused to carapace. 

Antennule (Fig. Ib): unsegmented with four aesthetascs/setae. 

Antenna (Fig. Ic): spinous process as long as rostral spine and with many subacute denticles; exopod very 

small, about one ninth of spinous process length. 

Maxillule (Fig. Id): endopod 2-segmented, proximal segment with one seta, distal with one subterminal and 

four terminal setae; basial endite with five spines, coxal with six spines/setae. 

Maxilla (Fig. le): endopod stepped distally, outer lobe broader than inner each with two long setae; basial 

endite incipiently bilobed distally, each with four setae; coxal endite bilobed distally, outer lobe with four and 

inner with five setae respectively, seta on apex of outer lobe very short almost a spine; scaphognathite with 

four plumose setae and a stout posterior process. 

First maxilliped (Fig. If): basis with eight setae arranged in pairs; endopod five-segmented with 1, 2, 1, 2, 4+ 1 

setae; exopod incipiently two-segmented and with four terminal plumose setae. 

Second maxilliped (Fig. Ig): basis with four setae; endopod three-segmented with 0, 1, 4+1 setae; exopod 

incipiently two-segmented with four terminal plumose setae. 

Third maxilliped and pereiopods: not developed. 

Abdomen (Fig. Ih): composed of five segments and a telson, somewhat dorso-ventrally compressed, surfaces 

with microscopic spinules; second and third segments with a pair of broad dorso-lateral processes; fourth 

segment laterally expanded and with a pair of obtuse lateral processes placed at a lower level than the ones on 

preceding segments; first segment, posterio-lateral margins truncate, those of other segments obtuse and of 

third to fifth with a very minute denticle; second to fifth segments each with a pair of setules near posterio- 

dorsal margin. Telson somewhat narrowed, furcae not noticeably directed outwards, each with numerous 

microscopic spinules and two very small lateral spinules; posterior margin with six equal plumose setae. 

Pachygrapsus transversus (Gibbes, 1850) 
Pachygrapsus transversus: Lebour, 1944: 115, fig. 5 (zoea I); Rossignol, 1957: 89, fig. 5 (megal.). 

Dimensions: T.T. 0-60-0-70 mm. C.L. 0-30-0-35 mm. Differs from P. gracilis in the following features. 
Carapace (Fig. 2a): dorsal spine longer and proximally slightly stouter, more than one third of carapace 
length; rostral spine noticeably stouter proximally; posterio-lateral margin of carapace with very 
inconspicuous denticles and with microscopic setules; only two pairs of anterio-median setules apparent. 
Antenna (Fig. 2c): spinous process with numerous subacute denticles developed distally as stout spine-like 
processes; exopod about one seventh of spinous process length. 
Maxillule (Fig. 2d): spines/setae slightly stouter. 
Maxilla (Fig. 2e): basial endite noticeably bilobed distally. 



34 



R. W. INGLE 




Fig. 1. Pachygrapsus gracilis (de Saussure). First zoea. a, carapace, right lateral aspect; b, antennule; c, 
antenna; d, maxillule; e, maxilla; f, first maxilliped; g, second maxilliped; h, abdomen and telson, 
dorsal aspect. Scale = 0-05 mm. 



FIRST ZOEA 



35 







Fig. 2. Pachygrapsus transversus (Gibbes). First zoea. a, carapace, right lateral aspect; b, antennule; c, 
antenna; d, maxillule; e, maxilla; f, first maxilliped; g, second maxilliped; h, abdomen and telson, 
dorsal aspect. Scale = 0-05 mm. 



36 R. W. INGLE 

Abdomen (Fig. 2h): slightly larger, posterio-lateral margins of fourth segment subacute and of the other 
segments (except first) more produced, denticles larger. Telson slightly broader, the more posterior of the two 
lateral spinules larger; furcae slightly shorter and stouter. 

Pachygrapsus marmoratus (Fabricius, 1787) 

Pachygrapsus marmoratus: Cano, 1892: S.Tav.III, figs IB (?zoea III), ? 1C, IE, IF, ?2c, 2e-f, ?3c, 4e-f, 5e-f, 
6e-f,7e-f,8e-f (labelled as 6 in fig.), 12e, 13e, 14e, 1 5e(?zoea IV, megal.); Williamson, 1915: 518, figs 403-^05, 
407-8 (figs after Cano); Hyman, 1924: 2, PI. 3, figs 22, ?23, 25, 26, ?33, 36a-b, 41, 42, 44, 45, 48, 49, 50, 52, 53, 
54, 56 (figs after Cano): Bourdillon-Casanova, 1960: 188, fig. 61 (zoea I); Paula, 1985: 142, fig. 3 (zoea I). 

Dimensions: T.T. 0-75 mm. C.L. 0-35 mm. 

Differs from P. gracilis and P. transversus as follows. 

Carapace (Fig. 3a): dorsal spine longer, exceeding half carapace length; posterio-lateral margin of carapace 

without setules, denticles very minute; anterio-median setules not apparent. 

Antenna (Fig. 3c): denticles on spinous process small and numerous throughout length of process. 

Maxillule (Fig. 3d): setules on spines/setae very long. 

Abdomen (Fig. 3h): fourth segment not laterally expanded and without a pair of obtuse lateral processes; 

posterio-lateral margins of third to fifth segments each with a conspicuous denticle. Telson lateral spinules on 

furcae absent. 

Family XANTHIDAE MacLeay, 1838 
Cataleptodiusfloridanus (Gibbes, 1850) 

Dimensions: T.T. 1-1 mm. C.L. 0-43 mm. 

Carapace (Fig. 4a): dorsal spine long, distally curved, proximally stout; rostral spine almost as long as dorsal 

spine and with 1 or 2 small spinules; lateral spines small; dorso-median elevation hardly developed; no 

anterio-median setules apparent, a small pair of posterio-median setules present; posterio-lateral margin of 

carapace with one or two small setules. 

Eyes: partly fused to carapace. 

Antennule (Fig. 4b): unsegmented and with four aesthetascs/setae. 

Antenna (Fig. 4c): spinous process as long as rostral spine, distally with many long acute spines; exopod small, 

between one sixth and one seventh of spinous process length and with two small distal setules. 

Maxillule (Fig. 4d): endopod two-segmented, proximal segment with one distal seta, distal segment with five 

setae (two subdistal and three distal); basial endite with five spines/setae; coxal endite with seven setae. 

Maxilla (Fig. 4e): endopod two-lobed, outer slightly stepped, broader than inner and with 2 + 2 setae, inner 

lobe with three setae; basial endite two-lobed, outer prominent and with four setae, inner with five setae; coxal 

endite two-lobed each with four setae; scaphognathite with four plumose setae and a stout posterior process. 

First maxilliped (Fig. 40: basis with ten setae arranged 2, 2, 3, 3, respectively; endopod five-segmented, with 

3, 2, 1, 2, 4+ 1 setae respectively; exopod incipiently two-segmented with four terminal plumose setae. 

Second maxilliped '(Fig. 4g): basis with four setae; endopod three-segmented, with 1,1,4+1 setae respectively; 

exopod incipiently two segmented, with four terminal plumose setae. 

Third maxilliped and pereiopods: not developed. 

Abdomen (Fig. 4h): composed of five segments and a telson; second segment with a pair of subacute laterally 

directed dorso-lateral processes, third segment with a pair of small acute posteriorly directed dorso-lateral 

processes; posterio-lateral margins of second segment acute, those of third to fifth segments extended into 

acute processes; posterio-dorsal surface of second to fifth segments each with a small pair of setules near 

margin; posterior margins of segments four and five with minute denticles. Telson furcae diverging slightly, 

each with one long prominent dorsal spine and one long and one smaller lateral spine, furcae with minute 

denticles; posteror margin of telson with six long setae. 



Remarks 

As mentioned earlier, some of the stages described and figured by Cano (1892) as Pachygrapsus 
marmoratus may not belong to this species. His figure depicting a first stage zoea (Tav.III, Fig. 1 A) 
does not show a dorso-lateral process on the third segment of the abdomen characteristic of 
Pachygrapsus zoeae and obvious in the present laboratory reared material. Cano's figure IB clearly 
shows this lateral process and although this larva is depicted with four maxillipedal exopod setae it 



FIRST ZOEA 



37 







Fig. 3. Pachygrapsus marmoratus (Fabricius). First zoea. a, carapace, right lateral aspect; b, antennule; 
c, antenna; d, maxillule; e, maxilla; f, first maxilliped; g, second maxilliped; h, abdomen and telson, 
dorsal aspect. Scale = 0-05 mm. 



38 



R. W. INGLE 




Fig. 4. Cataleptodius floridanus (Gibbes). First zoea. a, carapace, left lateral aspect; b, antennule; c, 
antenna; d, maxillule; e, maxilla; f, first maxilliped; g, second maxilliped; h, abdomen and telson dorsal 
aspect. Scale = 0-05 mm. 



FIRST ZOEA 39 

is probably of a later stage because he figures incipient pereiopods beneath the carapace. Hyman 
(1924) has suggested that this is a stage three zoea and was also convinced that Cano had over- 
looked a fourth stage and that the third stage described by Cano (purporting to be the last) 
represented the fifth and terminal zoeal stage of this species. Costlow & Bookhout (1962) however, 
maintained that . . . 'While some of Cano's (1891) figures may bear some slight inaccuracies, it is 
quite possible that the sequence and number of larval stages which he figures is correct. . .' These 
views were expressed in the context of their study of the larval development ofSesarma reticulatum 
in which there are only three zoeal stages and it is probable that P. marmoratus passes through five 
stages similar to P. crassipes. Cano (1892) also assigned two megalopal forms to P. marmoratus. 
The one illustrated in his fig. ID and bearing an acute rostral projection appears to be of an 
oxyrhynch as it lacks dactylar subterminal setae on the fifth pereiopods characteristic of brachy- 
rhynch megalopas. Bourdillon-Casanova (1960) described the first stage zoea of P. marmoratus 
from laboratory hatched material and illustrated (Fig. 61) prominent denticles on the carapace 
posterio-lateral margin as well as two conspicuous lateral setae (also shown by Paula, 1985, Fig. 
3,i) on each furca of the telson. These two features could not be detected in specimens examined 
during the present study. Lebour's (1944) figure of the first zoea of P. transversus, also obtained 
from Bermudan laboratory hatched crabs, differs from the present specimens in apparently having 
two distal setae on the antennal spinous process, a pair of conspicuous curved, acute dorso-lateral 
processes on the fourth abdominal segment and two equally developed lateral spinules on the 
telson furca. Her zoeae were also larger than the present ones. 

Mid-dorsal carapace setules have never been mentioned previously as occurring in Pachy- 
grapsus zoeae, and Gore & Scotto (1982: 518) suggested they may be absent in grapsinid zoeae. 
However in the present study these setules were found in first stage zoeae of P. gracilis and P. 
transversus but were not apparent in P. marmoratus. They are somewhat difficult to resolve 
satisfactorily, even with the aid of interference contrast, but appear to be less numerous on the 
anterio-median region of P. transversus than of P. gracilis. In many brachyuran larvae these setules 
do not appear until later stages. 

Martin (1984: 232-233) has provided an excellent key to the known xanthid zoeae of the Western 
Atlantic and Gulf of Mexico. The present study confirms his assessment of the larvae of this species 
as belonging to his group I of the xanthidae and they can be assigned to C. floridanus on the 
following combined features: (1) lateral processes. of third to fifth abdominal segments not 
extending posteriorly beyond half length of following segment, (2) more than twenty spinules on 
spinous process of antenna, (3) abdominal dorso-lateral processes confined to second and third 
segments, (4) telson with three spines on each furca, (5) lateral carapace spines present, (6) basial 
segment of first maxilliped endopod with three setae, (7) antennal exopod very reduced. 



Acknowledgements 

The visit to the Bermuda Biological Station was partly sponsored by an Exxon Corporation Fellowship. I 
thank Dr Wolfgang Sterrer, Director of the Biological Station and his staff for assistance during my visit 
and also Dr R. B. Manning for kindly presenting the P. marmoratus zoeae to the BM(NH). I also thank 
Dr A. L. Rice for reading the manuscript. 



References 

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Research Council. Tokyo. 2: 17-55. 
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Matematica e di Fisici della Societa Italiana della Scienze. Roma. (3) 8(4): 1-14. 
Costlow, J. D. Jr & Bookhout, C. G. 1962. The larval development of Sesarma reticulatum Say reared in the 

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40 R. W. INGLE 

Fabricius, J. C. 1 787. Mantissa insectorum sistens eorum species nuper detectas adjectis characteribusgenericis, 

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220-239. 
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Calcutta. 13(2): 154-165. 
Rathbun, M. J. 1923. The Brachyuran crabs collected by the U.S. Fisheries Steamer 'Albatross' in 1911, 

chiefly on the West Coast of Mexico. Bulletin of the American Museum of Natural History. New York. 

48(10): 6 19-637. 
Rice, A. L. 1980. Crab zoeal morphology and its bearing on the classification of the Brachyura. Transactions 

of the Zoological Society of London 35: 271424. 
Rice, A. L. & Williamson, D. 1. 1977. Planktonic stages of Crustacea Malacostraca from Atlantic Seamounts. 

Meteor Forschungsergebnisse. Berlin & Stuttgart. D26: 28-64. 
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Rossignol, M. & Roux, Ch. Mollusques, Crustaces, Poissons Marines des cotes d'A.E.F. en collection au 

Centre d'Oceanographie de 1'Institut d'Etudes Centrafricaines de Pointe-Noire. Ministere de la France 

d'Outre-Mer. Paris, pages 71-136. 
Saba, M. 1974. On the larval development of Chasmagnathus convexus de Haan (Grapsinae). Researches on 

Crustacea. The Carcinological Society of Japan. Tokyo. 6: 71-85. 
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Societe de Physique et d'Histoire Naturelle de Geneve 14: 41 7-496. 
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London 158: 357-370. 
Terada, M. 1981. Zoea larvae of five crabs in the subfamily Varuninae. Researches on Crustacea. The 

Carcinological Society of Japan. Tokyo. 11: 66-76. 



FIRST ZOEA 41 

Wear, R. G. 1970. Life-history studies on New Zealand Brachyura 4. Zoea larvae hatched from crabs of the 
family Grapsidae. New Zealand Journal of Marine and Freshwater Research. Wellington. 4(1): 3-35. 

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Manuscript accepted for publication 30 October 1985 



A classification of the phylum Sipuncula 

Peter E. Gibbs 

Marine Biological Association of the U.K., Plymouth, Devon PL1 2PB, U.K. 

Edward B. Cutler 

Division of Science and Mathematics, Utica College of Syracuse University, Utica, New York 
13502, U.S.A. 

Synopsis 

A classification of the phylum Sipuncula is adopted following the analysis of Cutler & Gibbs (1985) and 
comprises two classes, four orders and six families. This replaces the earlier classification of Stephen & 
Edmonds (1972) which was based on four families only. The diagnostic characters are reviewed. Seventeen 
genera are redefined, one new subgenus is described and twelve other subgenera are recognised. 

Introduction 

The classification of the phylum Sipuncula has had a confused history. Early attempts to define 
higher taxa by grouping genera were, to a large extent, thwarted by incomplete, imprecise or 
erroneous descriptions of many species. Stephen & Edmonds (1972) classified the phylum into 
four families in providing the first compilation of species described prior to about 1970. How- 
ever, this monograph is essentially literature-based and consequently many errors are repeated; 
nevertheless, it provides a useful base-line to the present revision. 

The need for greater precision in defining genera has led the authors to re-examine most of 
the available type specimens. The definitions of genera presented below incorporate both novel 
observations and corrections to earlier descriptions. Where possible, nine basic characters have 
been checked for each species before assigning it to a genus. These characters are summarised for 
each genus in Table 1 . A phylogenetic interpretation of the classification used here will be found in 
Cutler & Gibbs (1985). 



Diagnostic features of higher taxa 

In reviewing the diagnostic characters of the phylum, particular attention has been paid to the 
structure of the oral disk since the arrangement of the tentacles provides a useful basis for dividing 
the phylum into two classes - Sipunculidea and Phascolosomatidea. Certain descriptions of 
tentacle arrangements are misleading or in error (see for example Stephen & Edmonds (1972) p. 16 
and Table 3). No doubt these errors result from the fact that some species are not amenable to 
fixation in the extended state; species with long introverts are notoriously difficult to preserve with 
their introvert fully extended. Although dissection of the introvert is possible the details of the 
tentacular arrangement on a withdrawn disk are often difficult to interpret and have yet to be 
satisfactorily determined in some small-sized species (e.g. Apionsoma trichocephala Sluiter). The 
following summarises the distinctions of the two classes. 

The sipunculan tentacular crown exhibits many diverse forms but, basically, two tentacular 
patterns can be recognised. In one, that of the proposed class Sipunculidea, the tentacles are 
arranged peripherally on the oral disk so as to encircle the centrally-placed mouth; dorsally this 
circle is inflected to form an arc enclosing the nuchal organ, a feature well developed in Thysano- 
cardia spp for example (see Gibbs, Cutler & Cutler, 1983, Fig. 2). In the other, that of the proposed 
class Phascolosomatidea, the tentacles are restricted to a dorsal arc enclosing the nuchal organ and 



Bull. Br. Mus. not. Hist. (Zool.) 52(1): 43-58 Issued 29 January 1987 



44 



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Fig. 1. The structure of the tentacular crown of Sipuncula: some examples illustrating the form and 
variation within the classes Sipunculidea and Phascolosomatidea. Solid lines indicate possible evolu- 
tionary trends. A. Generalised Sipunculidea crown such as might have been possessed by ancestral 
stock adults; B. Golfingia margaritacea', C. Nephasoma rimicola; D. Onchnesoma squamatum; E. 
Nephasoma minutum; F. Sipunculus norvegicus', G. Themis te lageniformis; H. Phascolosoma 
granulatum; J. Antillesoma antillarum; K. Aspidosiphon johnstoni. (B, E, F: after Theel, 1905.) 
Abbreviations: cc, cephalic collar; m, mouth; n, nuchal organ; na, dorsal arc of tentacles enclosing 
nuchal organ; 1,2,3, primary, secondary and tertiary tentacle pairs around disk periphery. (Modified 
from Cutler & Gibbs, 1985.) 



there are no peripheral tentacles (Fig. 1). Thus, the two patterns have a common, perhaps homolo- 
gous, feature in the dorsal arc of tentacles. In evolutionary terms, the peripheral tentacles could be 
interpreted as a later addition, i.e. the Sipunculidea have evolved from a Phascolosomatidea stock. 
However, around the margin of the oral disk in Phascolosomatidea there is a prominent ridge, the 
cephalic collar, and it is thought that this ridge represents a vestige of the peripheral system. If 
this interpretation is correct the common ancestor must have possessed a Sipunculidea-type of 
tentacular crown, probably a simple form, somewhat similar to that of Thysanocardia procera 
(Gibbs, Cutler & Cutler, 1983, Fig. 2B), and the peripheral tentacles were lost during an early 
divergence to give the Phascolosomatidea line. The Sipunculidea tentacular pattern, peripheral 



46 P. E. GIBBS & E. B. CUTLER 

circle plus nuchal arc, can perhaps be best regarded as an elaboration of simple prostomial 
tentacles possessed by the early protostomial stock. 

The development of the tentacular crown in the Sipunculidea, as seen for example in Golfingia 
species, commences with the formation of four primary pairs of tentacles in the dorsal, ventral 
and lateral positions, between which secondary pairs subsequently develop to form a single ring 
encircling the central mouth on the oral disk (Fig. 1 A). Between these pairs tertiary pairs usually 
develop: in the adult these may be few or very numerous; in the latter case the tentacles are 
accommodated in loops or 'festoons' that extend aborally on to the anterior introvert. The nuchal 
organ situated dorsally between the two primary tentacles thus becomes enclosed by an arc of 
tentacles. As a general rule, the number of tentacles increases with increasing size and age of 
individuals and large-sized species have more tentacles than small-sized species. 

Within the class Sipunculidea a wide range of tentacular development is found. The most highly 
evolved crown is found in the genus Thysanocardia, adult specimens of which often possess 
well-developed festoons comprising several hundred tentacles; in some Thysanocardia nigra 
(Ikeda) the number exceeds 500 (see Gibbs, Cutler & Cutler, 1983). Large Golfingia margaritacea 
(Sars) have 100 or more tentacles (Fig. IB) but most other Sipunculidea have around 50 or fewer 
with only a limited number of tertiary tentacles developing, as in Golfingia elongata (Keferstein) 
with 20-34 and Nephasoma rimicola (Gibbs) with 12-20 (Fig. 1C; see Gibbs, 1973). In some species 
only the primary tentacles appear, as in Onchnesoma squamatum (Kor. & Dan.) with 8 and 
Nephasoma minutum (Keferstein) with just two (Fig. ID, E). Thus the evolution of the tentacular 
crown could have been not only towards greater complexity but also towards simplification, a 
trend, possibly neotenous, seen in several genera, notably Nephasoma (e.g. N. minutum), Onchne- 
soma (e.g. O. steenstrupi Kor. & Dan.) and Phascolion (e.g. P. pacificum Murina). Another vari- 
ation is seen in some members of the family Sipunculidae where the peripheral tentacles have 
become flattened and fused to form a continuous veil-like structure, as for example in Sipunculus 
norvegicus Dan. (Fig. IF). The crown of Themiste with its tentacles arising from 4-8 stems appears 
anomalous at first sight but, in fact, this type represents yet another modification of the basic 
Sipunculidea pattern. In themistids the secondary tentacles develop between the primary pairs but 
are borne on outgrowths of the oral disk so that with subsequent tertiary tentacle development, 
an erect dendritic structure results rather than the typical festoon which is contiguous with the 
introvert wall. In the themistid type the dorsal primary tentacles are widely spaced and do not 
enclose the nuchal organ (Fig. 1G). 

All six genera grouped in the class Phascolosomatidea are rather similar in terms of the tentacu- 
lar arrangement: with one exception, all have a single arc of up to 30 tentacles enclosing the nuchal 
organ (Fig. 1H,K). The exception is Antillesoma antillarum (Grube & Oersted) in which the 
tentacles are fairly numerous (Fig. 1 J) presumably as a result of secondary proliferation. 

One other character that separates the Sipunculidea and Phascolosomidea is the structure of the 
introvert hooks on the anterior introvert, when present. In the former group these hooks are 
somewhat variable but generally are simple, sharply-pointed protrusions of the epidermis and 
scattered in their distribution; however, in the latter they have a typical recurved shape, usually an 
internal structure is apparent and they are closely-packed in distinct rings encircling the anterior 
introvert. 

Definitions of orders, families and genera are given below. Four orders are recognised. In the 
class Sipunculidea, members of the order Sipunculiformes are distinguished by the presence of 
banding in the longitudinal muscle of the body wall found in five genera, all of which are placed in 
the family Sipunculidae. The remaining six genera within this class all have a uniform, continuous 
layer of longitudinal muscle tissue and form the order Golfingiiformes comprising three families 
Golfingiidae, Phascolionidae and Themistidae. In the class Phascolosomatidea the genera are 
separated into two orders, each with a single family, on the basis of the presence (Aspidosiphoni- 
formes: Aspidosiphonidae) or absence (Phascolosomatiformes: Phascolosomatidae) of an anal 
shield, a hardened thickening of the anterior trunk region. It should be noted that the structure of 
the anal shield is different in all three genera within the Aspidosiphonidae and it is recognised that 
this character may have evolved several times. 

Whilst the forms of the tentacle crown and of the introvert hooks are useful characters for 



SIPUNCULA 47 

dividing the 1 7 genera into two natural groups, here designated as classes, few other major charac- 
ters are confined to one or other of these two classes (Table 1): coelomic spaces in the body wall is a 
feature exclusive to Sipunculidae and likewise for anal shield development in Aspidosiphonidae. 
Other characters are found in both classes, notably the banding of the longitudinal muscle layer in 
the body wall, the attachment of the spindle muscle to the posterior trunk and an increase in the 
volume of the contractile vessel through the development of villi in conjunction with increased 
tentacular volume or area. Such characters would appear to be polyphyletic in origin. There is little 
doubt that the basic number of introvert retractor muscles is four, arranged as dorsal and ventral 
pairs. Loss of the dorsal pair appears to have occurred independently in a number of generic lines. 
Assessing the number of retractors in any one specimen can often be problematical because fusion 
frequently occurs but may not be evident. In some species only one retractor is apparent in the 
adult form: in Phascolion species there is good evidence to suggest the one muscle is the result of the 
fusion of all four muscles (Gibbs, 1985) whilst in Onchnesoma the single muscle is thought to 
comprise only the fused ventral pair, the dorsal pair having been lost. Use of the number of 
retractors as a taxonomic character has to be approached with some caution since even within a 
single population the number is liable to variation, as noted for Golfingia elongata (Gibbs, 1973). 
Morphological variation seems to be one of the hallmarks of the phylum, a feature that may 
account for the survival of this small group but one that does not facilitate good taxonomy. 

The present scheme of classification (Table 2) updates that given in Stephen & Edmonds (1972) 
and some later authors by incorporating the recent revisions of several major genera, notably, 
Siphonosoma, Golfingia and Phascolosoma. Synonymies are as given in Stephen & Edmonds 
(1972): any more recent changes are noted under each genus. 



Key to Families 

1 Tentacles arranged in an arc encircling dorsal nuchal organ; peripheral tentacles absent; hooks 

complex, in distinct rings [Class PHASCOLOSOMA TIDEA] 

Tentacles arranged peripherally on oral disk so as to encircle central mouth; may be borne 
on stem-like outgrowths of oral disk or reduced in number to a single dorsal pair; hooks simple, 
usually scattered [Class SIPUNCULIDEA] 

2 Anal shield present Fam. ASPIDOSIPHONIDAE (p. 55) 

Anal shield absent Fam. PHASCOLOSOMATIDAE(p. 54) 

3 Longitudinal muscles of body wall gathered into separate or anastomosing bands 

Fam. SIPUNCULIDAE (p. 48) 

Longitudinal muscle of body wall in a uniform continuous layer 

4 Tentacles carried on 4-8 stem-like outgrowths of oral disk. . Fam. THEMISTIDAE(p. 53) 
Tentacles not carried on disk outgrowths 

5 A single nephridium present Fam. PHASCOLIONIDAE(p. 51) 

Two nephridia present Fam. GOLFINGIIDAE(p. 50) 



Classification 

Phylum SIPUNCULA 
Class SIPUNCULIDEA 

Sipuncula with tentacles encircling a central mouth on the oral disk. Introvert hooks (when 
present) simple, thorn-like hollow structures that are usually irregularly distributed. Spindle 
muscle unattached posteriorly (except in Siphonosoma and Siphonomecus). 

Order SIPUNCULIFORMES 

Sipunculidea with longitudinal muscle in body wall gathered into bands (likewise for circular 
muscle in two genera Sipunculus and Xenosiphon). Coelomic extensions - canals or sacs - in body 
wall (except in Phascolopsis). 



48 P. E. GIBBS & E. B. CUTLER 

Table 2. Classification of the phylum Sipuncula 

Cl. Sipunculidea 
Ord. SIPUNCULIFORMES 

Fam. SIPUNCULIDAE Stephen & Edmonds, 1972 
Sipunculus Linnaeus, 1 766 
S. (Sipunculus) 

S. ( Austrosiphon ) Fisher, 1954 
Xenosiphon Fisher, 1947 
Siphonosoma Spengel, 1912 
Siphonomecus Fisher, 1 947 
Phascolopsis Fisher, 1950 
Ord. GOLFINGIIFORMES 

Fam. GOLFINGIIDAE Stephen & Edmonds, 1972 
Golfingia Lankester, 1885 
Nephasoma Pergament, 1946 
Thysanocardia Fisher, 1950 
Fam. PHASCOLIONIDAE Cutler & Gibbs, 1985 
PhascolionTheel 1875 
P. (Phascolion) 

P. (Isomya) Cutler & Cutler, 1985 
P. (Montuga) Gibbs, 1985 
P. (Lesenka) Gibbs, 1985 
P. (Villiophora) Cutler & Cutler, 1985 
Onchnesoma Koren & Danielssen, 1875 
Fam. THEMISTIDAE Cutler & Gibbs, 1985 
ThemisteGray, 1828 
T. (Themiste) 

T. (Lagenopsis) Edmonds, 1980 
Cl. Phascolosomatidea 
Ord. PHASCOLOSOMATIFORMES 

Fam. PHASCOLOSOMATIDAE Stephen & Edmonds, 1972 
Phascolosoma Leuckart, 1 828 
P. (Phascolosoma) 
P. ( Edmondsius ) subgen. nov. 
Apionsoma Sluiter, 1902 
Antillesoma Stephen & Edmonds, 1972 
Ord. ASPIDOSIPHONIFORMES 
Fam. ASPIDOSIPHONIDAE Baird, 1 868 
Aspidosiphon Diesing, 1851 
A. (Aspidosiphon) 
A. (Paraspidosiphon) Stephen, 1964 
Cloeosiphon Grube, 1 868 
Lithacrosiphon Shipley, 1902 



Family SIPUNCULIDAE Baird, 1868 
Characters are those of the order. 

Key to Genera 

1 Body wall circular muscle layer continuous PHASCOLOPSIS 

Body wall circular muscle layer gathered into bands 2 

2 Body wall circular and longitudinal muscle bands anastomosing, spindle muscle attached to 

posterior end of trunk 3 

Body wall circular and longitudinal muscle bands not anastomosing, spindle muscle not attached 
to posterior trunk 4 

3 Four introvert retractor muscles SIPHONOSOMA 

Two introvert retractor muscles . SIPHONOMECUS 



SIPUNCULA 49 

4 Gut with post-oesophageal loop; coelom extends into body wall as longitudinal canals running 
throughout most of trunk length ......... SIPUNCULUS 

Gut without post-oesophageal loop; coelom extends into body wall as short diagonal canals 
running across the width of one circular muscle band .... XENOSIPHON 

Genus SIPUNCULUS Linnaeus, 1766 

DIAGNOSIS. Introvert much shorter than trunk, without hooks, covered with scattered sub- 
triangular papillae. Trunk cylindrical. Body wall contains coelomic extensions in the form of 
parallel longitudinal canals which extend most of trunk length. Circular and longitudinal muscle 
layers gathered into distinct bands. Oral disk carries tentacles arranged around the mouth some- 
times modified with the development of an inter-tentacular membrane (S. nudus L., S. norvegicus 
Danielssen). Four introvert retractor muscles. Two protractor muscles may be developed (S. 
mundanus Sel. & Billow). Two contractile vessels, both without villi. Gut with post-oesophageal 
loop, caecum on rectum, and coil attached to body wall along its entire length by many connective 
strands. Spindle muscle not attached posteriorly. Two nephridia. Species usually large-sized (trunk 
greater than 5 cm long in adults). 

TYPE SPECIES. Sipunculus nudus Linnaeus, 1766, subsequent designation, ?Fisher 1952. 



Subgenus S/Pt/yVCt/LjyS Linnaeus, 1766 
Sipunculus (Sipunculus): Cutler & Cutler, 1985a: 232. 

DIAGNOSIS. Nephridia anterior to anus. Spindle muscle originates on body wall anterior to anus. 
TYPE SPECIES. Sipunculus nudus Linnaeus, 1766, subsequent designation, ?Fisher, 1952. 

Subgenus AUSTROSIPHON Fisher, 1954, emended 

Xenosiphon (Austrosiphon) Fisher, 1954: 314. 
Xenosiphon (Xenopsis) Johnson, 1969: 44. 
Sipunculus (Contraporus) Cutler & Cutler, 1985a: 241. 

DIAGNOSIS. Nephridia posterior to anus. Spindle muscle originates from ventral surface of rectum. 
TYPE SPECIES. Sipunculus mundanus Selenka & Biilow, 1883, monotypy. 

Genus XENOSIPHON Fisher, 1947 

DIAGNOSIS. Introvert much shorter than trunk, and without hooks but covered with scattered 
subtriangular papillae. Body wall contains coelomic extensions in form of short, diagonal canals 
limited in length to width of one circular muscle band. Circular and longitudinal muscle layers 
divided into distinct bands. Oral disk carries tentacles arranged around mouth. Four introvert 
retractor muscles and two thin protractor muscles present. Contractile vessel without villi, gut 
without post-oesophageal loop, caecum present on rectum and coil attached to body wall along 
entire length by connective strands. Spindle muscle originates on ventral wall of rectum and is not 
attached to the body wall posteriorly. Anus anterior to nephridiopores. Two nephridia. Contains 
one large-sized species. 

TYPE SPECIES. Sipunculus mundanus var. branchiatus Fischer, 1895, original designation. 

REMARKS. The two subgenera previously included in this genus are now assigned to Sipunculus (see 
above). 

Genus SIPHONOSOMA Spengel, 1912 

Siphonosoma (Siphonosoma): Fisher, 1950b: 805. 
Siphonosoma (Hesperosiphon) Fisher, 1950b: 805. 
Siphonosoma (Dasmosiphon) Fisher, 1950b: 805. 

DIAGNOSIS. Introvert much shorter than the trunk with prominent conical papillae (sometimes also 
hooks) arranged in rings. Body wall with coelomic sac-like extensions; circular and longitudinal 



50 P. E. GIBBS & E. B. CUTLER 

muscle layers gathered into anastomosing bands. Oral disk carries tentacles arranged around the 
mouth. Four introvert retractor muscles. Contractile vessel with or without villi. Spindle muscle 
attached posteriorly. Two nephridia. Species usually large-sized (trunk greater than 5 cm long in 
adults). 

TYPE SPECIES. Phascolosoma australe Keferstein, 1865, subsequent designation, Gerould, 1913. 

REMARKS. The three subgenera recognised by Fisher (1950b) were distinguished by the presence or 
absence of transverse dissepiments and rectal caeca. These characters have been found to be 
subject to great variation and of limited diagnostic value: consequently, this subgeneric separation 
is not supportable (Cutler & Cutler, 1982). 

Genus SIPHONOMECUS Fisher, 1947 

DIAGNOSIS. Introvert much shorter than trunk with prominent hooks and conical papillae 
arranged in rings. Body wall with coelomic extensions (sacs); circular and longitudinal muscle 
layers gathered into anastomosing bands. Oral disk carries tentacles arranged around the mouth. 
Two introvert retractor muscles. Contractile vessel without villi. Spindle muscle attached 
posteriorly. Two nephridia. Contains one large-sized species. 

TYPE SPECIES. Siphonomecus multicinctus Fisher, 1947, original designation. 

Genus PHASCOLOPSIS Fisher, 1950 

DIAGNOSIS. Introvert shorter than trunk with deciduous hooks (present in juvenile but lost in 
adult). Body wall without coelomic extensions. Circular muscle layer continuous, longitudinal 
muscle layer gathered into anastomosing bands. Oral disk carries tentacles arranged around the 
mouth. Four introvert retractor muscles. Contractile vessel without villi. Spindle muscle not 
attached posteriorly. Two nephridia. Contains one large-sized species. 

TYPE SPECIES. Sipunculus gouldii Portales, 1851, monotypy. 

Order GOLFINGIIFORMES 

Sipunculidea with body wall longitudinal muscle in a continuous layer, not gathered in bands. 

Family GOLFINGIIDAE Stephen & Edmonds, 1972 
Golfingiiformes with two nephridia. Tentacles not borne on stem-like extensions of oral disk. 

Key to Genera 

1 Contractile vessel with numerous villi THYSANOCARD1A 

Contractile vessel without villi 2 

2 Four introvert retractor muscles GOLFINGIA 

Two introvert retractor muscles NEPHASOMA 

Genus GOLFINGIA Lankester, 1885 

Golfingia (Golfingia): Fisher, 1950a; 549. 
Golfingia (Dushana) Murina, 1975: 1085. 
Themis te (Stephensonum) Edmonds, 1980: 33. 
Centrosiphon Shipley, 1903: 173. 

DIAGNOSIS. Introvert about equal to or shorter than trunk; hooks when present are usually scat- 
tered (arranged in rings in G. elongatd). Body wall with continuous muscle layers. Oral disk carries 
tentacles arranged around the mouth. Four introvert retractor muscles. Contractile vessel without 
villi. Spindle muscle not attached posteriorly. Two nephridia. Species small- to large-sized. 

TYPE SPECIES. Golfingia macintoshii Lankester, 1885 [ = Sipunculus vulgaris de Blainville, 1827: 
Stephen, 1934], monotypy. 



SIPUNCULA 5 1 

REMARKS. This genus now contains only those species previously assigned to the nominate sub- 
genus Golfingia (Golfingia). It includes Centrosiphon Shipley, 1903: Edmonds (1980) placed the 
type species C. herdmani Shipley within the genus Golfingia; the Centrosiphon specimens recorded 
by Cutler & Cutler (1979) are now considered to be aberrant Aspidosiphon. 

The subgenus G. (Dushana) Murina, 1975, was characterised by complete or partial fusion of 
the dorsal and ventral retractor muscles on one side of the body. However, it is known that such 
fusion of the retractors, and also reduction of the retractor number through loss of one or both 
dorsal retractors, are features of some Golfingia species, for example G. elongata (see Gibbs, 1973). 
The holotype of G. (Dushana) adriatica Murina shows a similar retractor arrangement (Murina, 
1975, Fig. 1) to that described by Watier (1932) for aberrant G. vulgaris. The type species originally 
designated for G. (Dushana), G. scutiger (Roule), does not differ significantly in its retractor 
arrangement (Roule, 1906, Fig. 95) from typical Golfingia species. Thus G. (Dushana) is no longer 
recognised. 

Genus NEPHASOMA Pergament, 1946 
Golfingia ( Phascoloides ) Fisher, 1950a: 550. 

DIAGNOSIS. Introvert about equal to, or shorter than, trunk. Hooks when present usually scattered 
(arranged in rings in N. rimicola (Gibbs), in spirals in TV. abyssorum (Kor. & Dan.)). Body wall with 
continuous muscle layers. Oral disk carries tentacles arranged around the mouth but tentacles may 
be reduced in both size and number and restricted to dorsal region. Two introvert retractor muscles 
often partially fused. Contractile vessel without villi. Spindle muscle not attached posteriorly. Two 
nephridia. Species generally small- to medium-sized (trunk less than 5 cm in length). 

TYPE SPECIES. Nephasoma marinki Pergament, 1946 [ = Onchnesoma glaciale Danielssen & Koren: 
Cutler & Murina, 1977; = Phascolosoma HlljeborgiiDsimdssen & Koren: Gibbs, 1982], monotypy. 

REMARKS. This genus now contains all those species previously assigned to the Golfingia subgenus 
Phascoloides Fisher, 1950, since Nephasoma Pergament has been shown to have priority over 
Phascoloides (Cutler & Murina, 1977). 

Genus THYSANOCARDIA Fisher, 1950 

DIAGNOSIS. Introvert longer than trunk, without hooks. Body wall with continuous muscle layers. 
Oral disk carries tentacles arranged around the mouth; those enclosing nuchal organ are well 
developed. Two introvert retractor muscles. Contractile vessel with distinct villi. Spindle muscle 
not attached posteriorly. Two nephridia. Species small- to medium-sized (adults generally under 
5 cm in trunk length). 

TYPE SPECIES. Phascolosoma procerum Mobius, 1875, original designation. 

REMARKS. The subgenus Golfingia (Thysanocardia) was recently elevated to generic rank and the 
number of species reduced to three by Gibbs, Cutler & Cutler (1983). 

Family PHASCOLIONIDAE Cutler & Gibbs, 1985 

Golfingiiformes with one nephridium (usually the right). Tentacles not borne on stem-like 
extensions of oral disk. Gut coil without well-defined axial spindle muscle. 

Key to Genera 

1 Anus usually situated on anterior trunk; epidermal 'holdfast' or 'attachment' papillae often 
present. Retractor muscles highly fused but usually 2-4 roots apparent at base of column 

PHASCOL1ON 

Anus situated on distal half of introvert; epidermal 'attachment' papillae absent. Retractor 
muscle(s) appear as single column without separate roots . . . ONCHNESOMA 



52 P. E. GIBBS & E. B. CUTLER 

Genus PHASCOLION Thcel, 1875 

DIAGNOSIS. Introvert length one-half to four times that of trunk length, with or without hooks. 
Trunk usually with modified 'holdfast' papillae. Body wall with continuous muscle layers. Oral 
disk carries tentacles arranged around the mouth. Introvert retractor muscle system modified by 
fusion of dorsal and ventral pairs: relative size and degree of fusion defines subgenera (see below). 
Contractile vessel without villi (but present in P. cirratum). Gut coiling generally loose and without 
axial spindle muscle. One nephridium (usually right). Species small- to medium-sized (less than 
5 cm in length) generally inhabiting mollusc shells. 

TYPE SPECIES. Sipunculus strombus Montagu, 1804, monotypy. 

Subgenus PHASCOLION Theel, 1875 
Phascolion (Phascolion): Gibbs, 1985: 314. 

DIAGNOSIS. Retractor column divided for most of its length: oesophagus detaches from retractor 
column at a point posterior to the first separation of the retractor muscles. Dorsal retractor(s) 
much more strongly developed than ventral retractor(s). Contractile vessel without villi. 

TYPE SPECIES. Sipunculus strombus Montagu, 1804, monotypy. 

Subgenus ISOMYA Cutler & Cutler, 1985 
Phascolion (Isomya) Cutler & Cutler 1985A: 820 

DIAGNOSIS. Characters as for P. (Phascolion) except that dorsal and ventral retractor muscles are 
about equal in diameter. 

TYPE SPECIES. Phascolion tuberculosum Theel, 1875, original designation. 

Subgenus MONTUGA Gibbs, 1985 
Phascolion (Montuga) Gibbs, 1985: 315. 

DIAGNOSIS. Retractor column divided only at posterior end: oesophagus detaches from retractor 
column at a point anterior to the first separation of the retractor muscles. Contractile vessel 
without villi. 

TYPE SPECIES. Phascolion Intense Selenka, 1885, original designation. 

Subgenus LESENKA Gibbs, 1985 
Phascolion (Lesenka) Gibbs, 1985: 315. 

DIAGNOSIS. Retractor column entire with retractor muscles fused throughout whole length. 
Contractile vessel without villi. 

TYPE SPECIES. Phascolion cryptum Hendrix, 1975, original designation. 

Subgenus V1LLIOPHORA Cutler & Cutler, 1985 
Phascolion (Villiophora) Cutler & Cutler, 19856: 821. 

DIAGNOSIS. Retractor column entire with retractor muscles fused throughout whole length. 
Contractile vessel with numerous villi. 

TYPE SPECIES. Phascolion cirratum Murina, 1968, monotypy. 

Genus ONCHNESOMA Koren & Danielssen, 1875 

DIAGNOSIS. Introvert much longer than trunk. Body wall with continuous muscle layers. Oral disk 
carries tentacles arranged around mouth but tentacles may be highly reduced in size. Introvert 
retractor muscle system modified by fusion to form single retractor muscle. Anus situated on 



SIPUNCULA 53 

introvert. Contractile vessel rarely apparent and without villi. Spindle muscle absent. One 
nephridium (right). Species small-sized (trunk less than 1 cm in length). 

TYPE SPECIES. Onchnesoma steenstrupii Koren & Danielssen, 1875, subsequent designation, 
Stephen & Edmonds, 1972. 

Family THEMISTIDAE Cutler & Gibbs, 1985 
Golfingiiformes with two nephridia. Tentacles borne on stem-like extensions of oral disk. 

Genus THEMISTE Gray, 1828 

DIAGNOSIS. Introvert less than trunk length. Body wall with continuous muscle layers. Oral disk 
carries tentacles basically surrounding mouth but extending with growth along margins of stem- 
like outgrowths of the oral disk. With or without hooks. Two introvert retractor muscles. Contrac- 
tile vessel with villi. Spindle muscle not attached posteriorly. Two nephridia. Species small- to 
large-sized. 

TYPE SPECIES. Themiste hennahi Gray, 1 824, monotypy . 

Subgenus THEMISTE Gray, 1828 
Themiste (Themiste): Edmonds, 1980: 33. 
DIAGNOSIS. Contractile vessel with long, thread-like villi. 
TYPE SPECIES. Themiste hennahi Gray, 1828, monotypy. 

Subgenus LAGENOPSIS Edmonds, 1980 
Themiste (Lagenopsis) Edmonds, 1980: 33. 
DIAGNOSIS. Contractile vessel with short, digitiform villi. 
TYPE SPECIES. Themiste lageniformis Baird, 1868, original designation. 

REMARKS. The subgenus T. (Stephensonum) Edmonds, 1980, was erected to include two species of 
Themiste having four, not two, retractor muscles, namely, Themiste stephensoni (the type species, 
original designation) and T. pinnifolia. The type material of Themiste stephensoni (Stephen) 
(described under the name Dendrostomum Grube, a junior synomym) in the RSME collections has 
been examined. The holotype (1958.23.24) has a golfingiid, not themistid, tentacle crown (as shown 
by dissection of the introvert) and the 'band of very short villi' on the contractile vessel (Stephen, 
1 942, p. 252) do not appear to be true villi but rather outpoutchings of a relatively voluminous 
vessel. The specimen is clearly a Golfingia and probably G. capensis (Teuscher); the other type 
specimens comprise further Golfingia but also include some Themiste all of which have the typical 
number of retractors (two). Thus T. (Stephensonum) becomes a junior synonym of Golfingia. The 
species Themiste pinnifolia (Keferstein) is based on a single specimen, collected more than 100 years 
ago, which cannot be traced. No subsequent record appears in the literature, despite extensive 
collecting in the area of the type locality (St Thomas, West Indies). Since the generic identity of this 
specimen is in doubt, the species name pinnifolia is regarded as a nomen dubium. 

Class PHASCOLOSOMATIDEA 

Sipuncula with tentacles confined to an arc enclosing dorsal nuchal organ: peripheral tentacles 
absent. Introvert hooks recurved, usually with an internal structure and closely-packed in 
regularly-spaced rings (absent in Antillesoma}. Spindle muscle attached posteriorly. 

Order PHASCOLOSOMATIFORMES 

Phascolosomatidea with anterior trunk not modified to form anal shield. Four introvert retractor 
muscles. 



54 P. E. GIBBS & E. B. CUTLER 

Family PHASCOLOSOMATIDAE Stephen & Edmonds, 1972 
Characters are those of the order. 

Key to Genera 

1 Introvert hooks absent. Contractile vessel with villi ANTILLESOMA 

Introvert hooks present. Contractile vessel without villi 2 

2 Longitudinal muscle in body wall gathered into bands .... PHASCOLOSOMA 
Longitudinal muscle in body wall a uniform continuous layer . . . . AP1ONSOMA 

Genus PHASCOLOSOMA Leuckart, 1828 

DIAGNOSIS. Introvert variable in length, often equal to trunk with numerous rings of recurved 
hooks (absent in P. meteori Herubel). Body wall with longitudinal muscle layer gathered into 
bands. Oral disk carries relatively few tentacles (less than 30) enclosing nuchal organ. Contractile 
vessel without true villi (may have bulbous vesicles). Four introvert retractor muscles; lateral pairs 
sometimes partially, rarely completely, fused. Spindle muscle attached posteriorly (except in 
P.pectinatwri). Two nephridia. 

TYPE SPECIES. Phascolosoma granulatum Leuckart, 1828, monotypy. 

Subgenus PHASCOLOSOMA Leuckart, 1828 

Phascolosoma (Phascolosoma): Stephen & Edmonds, 1972: 289. 
? Phascolosoma (Rueppellisoma) Stephen & Edmonds, 1972: 271. 
?Phascolosoma (Satonus) Stephen & Edmonds, 1972: 28 (in part). 

DIAGNOSIS. Spindle muscle attached posteriorly. Introvert hook without accessory spinelets. 
TYPE SPECIES. Phascolosoma granulatum Leuckart, 1828, monotypy. 

Subgenus EDMONDSIUS subgen. nov. 
Phascolosoma (Satonus) Stephen & Edmonds, 1972: 282 (in part) 

DIAGNOSIS. Spindle muscle not attached posteriorly. Introvert hook with accessory spinelets at 
base. 

TYPE SPECIES. Phascolosoma pectinatum Keferstein, 1867, monotypy. 
The subgenus is named in honour of Dr Stanley J. Edmonds. 

REMARKS. Stephen & Edmonds (1972) attempted to divide this large genus by creating four sub- 
genera, P. (Phascolosoma), P. (Rueppellisoma), P. ( Antillesoma) and P. (Satonus), for the most 
part using published descriptions concerning the number of retractor muscles (four or two), 
presence or absence of contractile vessel villi and whether or not the spindle muscle is attached 
posteriorly. In examining all of the available type material, Cutler & Cutler (1983) found that the 
subgeneric distinctions were highly confused because many of the original descriptions contained 
errors. P. (Rueppellisoma), comprising eight putative species each allegedly with two retractor 
muscles, is now considered invalid (all Phascolosoma are now interpreted as having four retrac- 
tors), the type species, Phascolosoma rueppellii Grube, 1868, by original designation, being placed 
as incertae sedis since the type is lost. P. (Antillesoma), formerly containing six species, now 
contains only the type species, Phascolosoma antillarum Grube & Oersted, 1858, original desig- 
nation; this subgenus is sufficiently distinct as to warrant generic rank (see below). The remaining 
subgenus, P. (Satonus), is distinguished from the nominate subgenus by the absence of a posterior 
attachment of the spindle muscle. This character is difficult to determine with any degree of 
certainty in any specimen that has been damaged internally, become macerated or has dried, as 
found when most of the type materials of the eight species grouped in P. (Satonus) were re- 
examined, including that of the type species, Phymosoma nigritorquatum Sluiter, 1 882, original 
designation. Just one species, Phascolosoma pectinatum Keferstein, 1867, appears to fit the defi- 
nition off. (Satonus). Since the type species of this subgenus, P. nigritorquatum, has uncertain 



SIPUNCULA 55 

status (it may be a junior synonym of P. ( Phascolosoma) scolops (Selenka & de Man)), it has been 
categorised as incertae sedis (Cutler & Cutler, 1983). Thus P. (Satonus) is invalid and the new 
subgenus accommodates P. pectinatum. 

Genus APIONSOMA Sluiter, 1902 

Apionsoma Sluiter, 1902:42. 

Golfingia (Mitosiphon) Fisher, 1950a: 550. 

Golfingia (Phascolana) Wesenberg-Lund, 1959: 183. 

Fisher ana Stephen, 1964: 460. 

Golfingia (Siphonoides) Murina, 1967: 1334. 

DIAGNOSIS. Introvert of variable length in relation to trunk with rings of recurved hooks (absent in 
A. trichocephald) that in some species have accessory spinelets at base. Body wall with continuous 
muscle layers. Oral disk with tentacles enclosing nuchal organ but not mouth. Contractile vessel 
without villi. Four introvert retractor muscles. Spindle muscle attached posteriorly. Two 
nephridia, sometimes bilobed. Species small-sized (less than 2 cm in length). 

TYPE SPECIES. Apionsoma trichocephala Sluiter, 1902, monotypy. 

REMARKS. Cutler (1979) reviewed this taxon which is here elevated to generic status. It includes 
many species previously assigned to various Golfingia subgenera and Fisher ana (see above). This 
genus is one that still presents problems, in particular, the precise nature of the oral disk in A. 
trichocephala remains unknown. The variations within the genus may justify the use of subgenera. 

Genus ANT1LLESOMA Stephen & Edmonds, 1972 
Phascolosoma (Antillesoma) Stephen & Edmonds, 1972: 277. 

DIAGNOSIS. Introvert variable in length, often about equal to trunk, without hooks. Body wall with 
longitudinal muscle layer gathered into anastomosing bands. Oral disk carries numerous tentacles 
(more than 30 in adults) enclosing nuchal organ. Contractile vessel with many villi. Four introvert 
retractor muscles, lateral pairs often extensively fused. Spindle muscle attached posteriorly. Two 
nephridia. Contains one small- to medium-sized species (less than 5 cm in length). 

TYPE SPECIES. Phascolosoma antillarum Grube & Oersted, 1858, original designation. 

REMARKS. This taxon was erected as a subgenus to include six Phascolosoma species but is now 
considered to be monospecific (Cutler & Cutler, 1983) and of generic rank. 

Order ASPIDOSIPHONIFORMES 

Phascolosomatidea with the anterior trunk hardened to form a horny or calcareous anal shield. 
Two retractor muscles. 

Family ASPIDOSIPHONIDAE Baird, 1868 
Characters are those of the order. 

Key to Genera 

1 Introvert protrudes from centre of anal shield. Shield calcareous (white) composed of numerous 

polygonal plates . CLOEOS1PHON 

Introvert protrudes from ventral margin of anal shield ... . .2 

2 Shield composed of single calcareous cap .... . LITHACROSIPHON 
Shield composed of numerous horny (brown-black) plates. . . . ASPIDOSIPHON 

Genus ASPIDOSIPHON Dieting, 1851 

DIAGNOSIS. Introvert usually longer than trunk with recurved hooks in numerous rings. Trunk 
with anal shield composed of hardened plates (occasionally inconspicuously developed). Introvert 



56 P. E. GIBBS & E. B. CUTLER 

protrudes from ventral margin of shield. Body wall either with continuous longitudinal muscle 
layer or with longitudinal muscle layer gathered into anastomosing, sometimes ill-defined, 
bands. Oral disk with tentacles enclosing nuchal organ but not mouth. Contractile vessel without 
villi. Two introvert retractor muscles often almost completely fused. Spindle muscle attached 
posteriorly. Two nephridia. Species small- to medium-sized. 

TYPE SPECIES. Aspidosiphon muelleri Diesing, 1851, subsequent designation, Stephen & Edmonds, 
1972. 

Subgenus ASPIDOSIPHON Diesing, 1851 
Aspidosiphon (Aspidosiphon): Cutler, 1973: 174. 
DIAGNOSIS. Longitudinal muscle layer of body wall continuous, not gathered into bands. 

TYPE SPECIES. Aspidosiphon muelleri Diesing, 1851, subsequent designation, Stephen & Edmonds, 
1972. 

Subgenus PARASPWOSIPHON Stephen, 1964 

Paraspidosiphon Stephen, 1964: 459. 
Aspidosiphon (Paraspidosiphon): Cutler, 1973: 168. 

DIAGNOSIS. Longitudinal muscle layer of body wall gathered into bands. 
TYPE SPECIES. Aspidosiphon steenstrupii Diesing, 1859, original designation. 

REMARKS. Earlier diagnoses of this genus contain serious errors. The tentacular arrangement is 
phascolosomatid (Gibbs, 1977; Gibbs, in Edmonds, 1980) and there are always two retractor 
muscles. Although A. semperi ten Broeke and A. insular is Lanchester are described as having four 
retractor muscles, the type of the former (ZMUA collection) has, in fact, two, and the type of the 
latter (BMNH: Reg. 1 924.3. 1 .80) is not an Aspidosiphon but a Phascolosoma (possibly P. perlucens 
Baird). In Aspidosiphon species the spindle muscle is always attached posteriorly. 

Genus CLOEOSIPHON Grube, 1868 

DIAGNOSIS. Introvert longer than trunk with numerous rings of recurved hooks. Trunk with 
conspicuous anal shield composed of small rectangular calcareous plates. Introvert protrudes 
through centre of shield. Body wall with continuous muscle layers. Oral disk carries tentacles 
enclosing nuchal organ, but not mouth. Contractile vessel without villi. Two introvert retractor 
muscles often almost completely fused. Spindle muscle attached posteriorly. Two nephridia. 
Contains one medium-sized species. 

TYPE SPECIES. Loxosiphon aspergillus Quatrefages, 1865, monotypy. 

Genus L1THACROSIPHON Shipley, 1902 

DIAGNOSIS. Introvert about equal to trunk with numerous rings of recurved hooks. Trunk with 
anal shield formed by internal calcareous conical structure. Body wall with longitudinal muscle 
layer gathered into bands. Oral disk with tentacles enclosing nuchal organ but not mouth. 
Contractile vessel without villi. Two introvert retractor muscles, often almost completely fused. 
Spindle muscle attached posteriorly. Two nephridia. Species small- to medium-sized (less than 
4 cm in length). 

TYPE SPECIES. Lithacrosiphon maldiviense Shipley, 1902, monotypy. 
REMARKS. This genus now contains two species (see Cutler & Cutler, 1981). 

Acknowledgements 

We are indebted to numerous colleagues for generously providing specimens and for kindly arranging loans 



SIPUNCULA 57 

of type materials, in particular, Mr R. W. Sims -British Museum (Natural History), Dr S. van der Spoel- 
Zoologisch Museum, Universiteit van Amsterdam, Dr R. Olerod - Naturhistoriska Riksmuseet, Stockholm, 
Dr J. B. Kirkegaard - Zoologisk Museum, Copenhagen, Dr C. B. Goodhart - University Museum of 
Zoology, Cambridge, and Dr S. Chambers - Royal Scottish Museum, Edinburgh. We are grateful to Mr 
R. W. Sims for his help with the preparation of this paper. 

References 

Cutler, E. B. 1973. Sipuncula of the western North Atlantic. Bulletin of the American Museum of Natural 

History 152: 103-204. 
1979. A reconsideration of the sipunculan taxa Fisherana Stephen, Mitosiphon Fisher and Apionsoma 

Sluiter. ZoologicalJournal of the Linnean Society 65: 367-384. 
& Cutler, N. J. 1979. Madagascar and Indian Ocean Sipuncula. Bulletin du Museum d'Histoire naturelle, 

Paris (Ser 4), 1:941-990. 
& 1981. A reconsideration of Sipuncula named by I. Ikeda and H. Sato. Publications of the Seto 

Marine Biological Laboratory 26: 51-93. 
& 1982. A revision of the genus Siphonosoma (Sipuncula). Proceedings of the Biological Society of 

Washington 95: 748-762. 
& 1983. An examination of the Phascolosoma subgenera Antillesoma, Rueppellisoma and Satonus. 

Zoological Journal of the Linnean Society 77: 175-187. 
& 1985a. A revision of the genera Sipunculus and Xenosiphon (Sipuncula). ZoologicalJournal of the 

Linnean Society 85: 219-246. 
& 19856. A revision of the genera Phascolion Theel and Onchnesoma Koren and Danielssen 

(Sipuncula). Proceedings of the Biological Society of Washington 98: 809-850. 
& Gibbs, P. E. 1985. A phylogenetic analysis of higher taxa in the phylum Sipuncula. Systematic Zoology 

34: 162-173. 

& Murina, V. V. 1977. On the sipunculan genus Golfingia Lankester, 1885. Zoological Journal of the 



Linnean Society 60: 173-187. 
Edmonds, S. J. 1980. A revision of the systematics of Australian sipunculans (Sipuncula). Records of the South 

Australian Museum 18: 1-74. 
Fisher, W. K. 1950a. The sipunculid genus Phascolosoma. Annals and Magazine of Natural History (Ser. 12) 3: 

547-552. 
1950b. Two new subgenera and a new species of Siphonosoma (Sipunculoidea). Annals and Magazine of 

Natural History (Ser 12), 3: 805-808. 
1952. The sipunculid worms of California and Baja California. Proceedings of the United States National 

Museum 102: 371-450. 

1954. The genus Xenosiphon (Sipunculoidea). Annals and Magazine of Natural History (Ser. 12), 7: 



311-315. 
Gerould, J. H. 1913. The sipunculids of the eastern coast of North America. Proceedings of the United States 

National Museum 44: 373-437. 
Gibbs, P. E. 1973. On the genus Golfingia (Sipuncula) in the Plymouth area with a description of a new species. 

Journal of the Marine Biological Association of the United Kingdom 53: 73-86. 

1977. British sipunculans. Synopsis of the British Fauna (New Series) 12, 35 pp. 

1982. The synonymy of the Golfingia species assigned to the abyssorum section (Sipuncula). Sarsia 67: 

119-122. 
1985. On the genus Phascolion (Sipuncula) with particular reference to the north-east Atlantic species. 

Journal of the Marine Biological Association of the United Kingdom 65: 3 1 1-323. 

Cutler, E. B. & Cutler, N. J. 1983. A review of the sipunculan genus Thysanocardia Fisher. Zoologica 



Scripta 12: 295-304. 
Johnson, P. 1969. A new subgenus of Xenosiphon (Sipunculidae) and description of a new species from Indian 

waters. Journal of the Bombay Natural History Society 66: 43^46. 
Murina, V. V. 1967. Report of the sipunculid worms from the sub-littoral zone of Cuba and the Mexican Gulf. 

Zoologicheskii Zhurnal 46: 1329-1339. [In Russian with English summary.] 
1975. New taxa of the genus Golfingia. Zoologicheskii Zhurnal 54: 1085-1089. [In Russian with English 

summary.] 
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annees 1880-1883, 8: 1-102. 
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Government of Ceylon on the Pearl Oyster Fisheries of the GulfofManaar. Part 1, Suppl. Rep. 3: 171-176. 



58 P. E. GIBBS & E. B. CUTLER 

Sluiter, C. Ph. 1902. Die Sipunculiden und Echiuriden der Siboga-Expedition. Siboga-Expeditie 25: 1-53. 
Stephen, A. C. 1934. The Echiuridae, Sipunculidae, and Priapulidae of Scottish and adjacent waters. 

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sipunculids of Cape Province and Natal. Annals of the Natal Museum 10: 245-256. 
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& Edmonds, S. J. 1972. The phyla Sipuncula and Echiura. 528 pp. London. British Museum (Natural 



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Manuscript accepted for publication 6 December 1985 



Two new species of Garra (Teleostei-Cyprinidae) 
from the Arabian peninsula 

K. . Banister 

Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD 



Introduction 

Continued interest in speleology in Oman has led to the discovery of yet another hypogean, but in 
this case microphthalmic, species of the cyprinid genus Garra from an extremely isolated sink hole 
in the Jabal Qara mountains of Oman. In the course of comparing this new species with its 
congeners, using the recent revision of Krupp (1983), the discovery of more specimens of Krupp's 
' Garra: incertae sedis (1)' enabled a second new species to be described. In all, eight species of Garra 
are now known from the Arabian peninsula. 



Garra dun sire i sp. nov. 
(Fig. 1) 

The first four specimens received (BMNH 1984.3.6: 577-580) were collected by Mr A. Dunsireand 
Mr D. Green on 16 May 1 980, but were in too poor a condition to be used for description. A further 
consignment of 19 live specimens (13 of which were still alive at the time of writing, July 1985) was 
collected by Mr D. Maclelland on 26 February 1983. 

HOLOTYPE. BMNH 1984.3.6: 571, 68 mm SL; Tawi Atair (1706'N, 5434'E) in the Jabal Qara 
(variously Jabal Samhan) mountains, Dhofar, Oman. 

PARATYPES. BMNH 1984.3.6: 572-576, 34-^9 mm, same data as holotype. 

LOCALITY. The fishes were caught in a pool in a side passage 200 m down the sink hole shown in 
Plates 1 and 2. The surface drainage is southwards to the Arabian Sea. The significance of the 
drainage direction and the isolated of the locality will be discussed below. 

DESCRIPTION. The description is based on the holotype and five paratypes (34-68 mm SL), all of 
which were radiographed. Additionally two of the first four specimens (1984.3.6: 577-580) were 
cleared and stained with alizarin. All measurements are expressed as a percentage of the standard 
length. Because the sample is so small, the measurements of the holotype are included in the range 
as well as being given in parentheses. 

MORPHOMETRIC DATA. Body depth x = 22-9, range = 21-4-23-8 (23-8); head length, x = 27-3, range = 25-0-30-3 
(25-0); eye diameter x = 3-3, range = 2-8-3-7 (3-7); interorbital width x = 9-4, range = 8-2-10-0 (8-8); pectoral 
fin length x = 20-2, range= 18-8-21-4 (19-3); caudal peduncle length x = 13-7, range= 12-0-16-0 (14-7); caudal 
peduncle depth x = 10-1, range = 9-2-1 1-4 (10-3); anterior barbel length x = 4-2, range = 3-7-4-5 (3-7); poster- 
ior barbel length x = 3-5, range = 2-2^-0 (2-6); dorsal fin height x = 23- 1 , range = 22-2-23-8 (22-8); mental disc 
maximum length x = 7-4, range = 6-5-8-8 (6-5); mental disc maximum width x = 7-3, range = 7- 1-8-1 (7-2); 
distance between snout and anus x = 76-6, range = 73-5-78-5 (74-4); distance between snout and anal fin origin 
x = 80-9, range = 77-9-83-6 (78-7); distance between snout and pelvic fin origin x = 60-5, range = 58 -8-63 -3 
(59-1); distance between snout and dorsal fin origin x = 53-0, range = 45-9-55-9 (51-5). 

The body shape and details of the mental disc are shown in Figs 1 and 2a. The abdomen of the 
holotype has collapsed, creating an uncharacteristic flat-bellied appearance. The eyes are very 
small and not visible in ventral view. The mental disc is approximately circular. In the smallest 
specimen (34 mm SL), only the posterior margin of the disc is free, but as the fish grows, the rest of 



Bull. Br. Mm. nat. Hist. (Zool.) 52(1): 59-70 Issued 29 January 1987 



60 



K. E. BANISTER 




Plate 1 . The sink hole at Tawi Atair . 



SPECIES OF GARRA 



61 




Plate 2. A close-up of the arrowed area in Plate 1 to indicate the size of the sink hole. 




Fig. 1. Garra dunsirei Holotype. Scale = 1 mm. 



62 



K. E. BANISTER 




Fig. 2. Ventral views of the heads of a. Garra dunsirei and b. Garra lautior. Scale = 5 mm. 



the rim becomes free. The papillae are scarcely developed, the papillar bed being only just differen- 
tiated in a specimen of 50 mm SL (Fig. 2a). Only on the holotype are the papillae well formed. The 
size and extent of the papillae are less than in all other Garra in the region. 

None of the specimens, alive or preserved, has tubercles on the snout, although a 39-5 mm SL 
specimen is a gravid female. 

SQUAMATION. The scales are less well developed than those of the hypogean population of Garra 
barreimiae (Banister, 1 984) which, although thinner than the scales of epigean populations, are 
scarcely less extensive. In Garra dunsirei the scales are as deep as, or deeper, than long and although 
they fill the scale pocket vertically, they often fail to reach the posterior edge of the pocket. A scale 
and its pattern of striations is shown in Fig. 3. Scales of the holotype have eleven 'growth' rings. It is 
not known if these represent annual or seasonal spawning marks or are the result of growth 
changes caused by food availability. Food only comes into the cave .during the annual rains. The 
scales of the ventral region of the body are severely reduced or absent. In the lateral line series there 
are 34(f2), 35(f3) or 36(fl) scales. There are 3^(f6) scale rows from the dorsal fin base to the lateral 
line and 2^(f3) or 3|(f3) scale rows from there to the pelvic fin base. On the two specimens on which 
a count was possible there are 6| scale rows from the lateral line to the ventral mid-line. Twelve (f6) 
scale rows encircle the least circumference of the caudal peduncle. 

VERTEBRAE. Radiographs revealed the presence of 32(f3) or 33(f3) vertebrae, excluding those 
forming the Weberian mechanism. It was difficult to identify with certainty the first caudal vertebra 
but there appear to be 18(f2)or 1 9(f4) abdominal vertebrae. There are 14(f2)or 15(f4) pairs of ribs. 
Characteristic of this species is the most unusual change of shape of the neural arches and spines 
below the dorsal fin. In all the other Arabian peninsular Garra species and in all other Garra species 
investigated the neural spines are shorter in this region, but of the same general configuration and 
angle as the other neural spines (Fig. 9). In Garra dunsirei (Fig. 4) the neural spines are bent sharply 
back and come to lie almost in parallel with the axis of the centrum. 



SPECIES OF GARRA 



63 




Fig. 3. A scale from the row above the lateral line of the holotype of Garra dunsirei. Scale = 0-5 mm. 




Fig. 4. Garra dunsirei. The vertebral column below the dorsal fin to show the unusual profile of the 
neural spines. For clarity the ribs are omitted. Scale = 1 mm. 



FINS. The dorsal fin has only 3(f6) unbranched and 7(f5) or 8(fl) branched rays. The foremost 
unbranched ray is minute and only visible in a radiograph or an alizarin preparation. The anal fin 
has 3 unbranched and 5 branched rays (f6). The first unbranched ray is again minute and not visible 
externally. 

GILL RAKERS. The gill rakers are small in size and few in number. There are 6(f2) or 7(f2) on the 
lower limb of the first gill arch. They could not be counted in the two smallest specimens. 

PHARYNGEAL BONES AND TEETH. The pharyngeal teeth number 2.4.5-5.4.2 (Fig. 5). The five 
posterior teeth of the innermost row are thin and have hooked crowns, quite unlike those typical of 
other Garra species from the Arabian peninsula (see Krupp, 1983: figs 23, 27, 40). 



64 K. E. BANISTER 




Fig. 5. Left pharyngeal bone of Garra dunsirei. Scale = 1 mm. 



COLORATION. Alcohol preserved specimens are uniform pale, yellowish grey. The ventral surface is 
only slightly paler than the rest of the body. There are traces of dark pigment near the anterior edge 
of the pectoral fins, on the dorsal edge of the dorsal fin membrane as well as near the base of the 
dorsal fin (the last being the typical Garra markings). Other fins are colourless. 

Living fishes are a dirty white, slightly more heavily dusted with dark pigment dorsally. The 
cheeks and operculum reflect greenish-gold. The post-opercular spot is inconspicuous but the red 
of the blood in the gills is clearly visible through the adjacent notch. 

ETYMOLOGY. This species is named in honour of the collector, Mr Andy Dunsire, who has 
encouraged so many people to search for subterranean fishes, as well as collecting such fish himself 
in isolated and hazardous regions. 

NOTES ON LIVING SPECIMENS. The fishes swim slowly, but continually, usually in a slightly head 
down position. When resting, they are indifferent to their orientation provided that their ventral 
surface is in contact with a solid object. They stay near the bottom of the aquarium, very rarely 
approaching the surface even to take food. 

In these aspects they contrast markedly with the blind, hypogean population of Garra barrei- 
miae which swims rapidly and swarms at the surface on the introduction of food (Banister, 1984). 

Garra dunsirei shows no preference for either light or dark conditions, although a strong light 
beam shone on an individual will cause it to jerk away after one or two seconds. After that, 
however, the fishes will come and investigate a localised illuminated patch on the substrate. 

DISTRIBUTION AND RELATIONSHIPS. Of particular interest is the geographical isolation of Garra 
dunsirei horn its congeners. In fact, no primary freshwater fishes have been recorded from this area 
of Dhofar, the closest natural populations being nearly 400 miles away at Tarim in the Wadi 
Hadramut to the west and also some 450 miles to the northeast in the Omani Jabal Akhdar. The 
drainage of this part of the peninsular coast is a series of isolated wadis carrying the run-off due 
south from the Jabal Qara range to the Arabian sea. On the north side of the Jabal Qara is an 
interlinked series of wadis that in wetter times would have carried water northwards into the 
extensive lake or lake system that occupied the site of the Rub al Khali. Krupp (1983) points out 
that the Jabal Qara range is part of an ancient feature whose presence caused the formation of the 
internal drainages. Although the internal basin would have permitted fish dispersal northward 
from Jabal Qara, whether or not it allows fishes from the north access to the streams south of Jabal 
Qara is unknown. It might be possible to determine the hydrological affinities of this isolated water 
source were the phylogenetic relationships of Garra dunsirei known. For the moment, therefore, 
neither the hydrological affinities of the sink hole nor the phylogenetic relationships of Garra 
dunsirei can be determined. 

DIAGNOSIS. This species can be characterised by the late development of the papillar beds on 
the mental disc, the papillar bed being only just differentiated at 50 mm SL; also the small eye 



SPECIES OF GARRA 



65 



diameter (x = 3-3) and the highly unusual shape of the neural spines below the dorsal fin (see Fig. 4 
and p. 62). 



Garra lautior sp. nov. 

The recognition of this species stems from Krupp (1983: 615) who described six specimens from the 
Wadi Hadramut as 'Garra: incertae sedis (1)' but was reluctant to base a species on such a small 
sample. A search through the collections of the British Museum (Natural History) revealed 13 
more specimens. 

HOLOTYPE. BMNH 1976.4.7: 398, 74 mm SL from the Qasam area, Wadi Hadramut, Yemen, coll. 
King-Webster. 

PARATYPES. BMNH 1976.4.7: 399-404, 64-74 mm SL (other details as above); 1976.4.7: 647-648, 
71 & 75 mm SL (other detals as above); 1976.4.7: 377-378, 61 & 74 mm SL: (other details as above); 
1976.4.7: 645, 80 mm SL from Al-Ghurf, Wadi Hadramut, coll. King- Webster; BMNH 1976.4.7: 
366 79 mm SL, from Gheil Umar, Wadi Hadramut, coll. King- Webster. 

DESCRIPTION. The description is based on the holotype as 12 paratypes (61-80 mm SL). The 
measurements are expressed as a percentage of the standard length those of the holotype are 
included in the range also also given separately in parentheses. 

MORPHOMETRIC DATA. Body depth x = 22-3, range = 20-2-25-0 (22-3) (n = 8); head length x = 22-4, 
range = 21 -6-23-4 (21-6); eye diameter x = 4-8, range = 4-0-5-4 (5-4); mouth width x = 5-7, range = 4-4-6-6 
(6- 1); pectoral fin length x= 19-4, range= 18-7-21 -5 (19-3); caudal peduncle length x= 17-1, range =15-2-20-0 
(15-9); caudal peduncle depth x = 8-6, range = 7-7-9-4 (9-3); anterior barbel length x = 2-6, range = 1-3-3-2 
(3-0); posterior barbel length x = 2-0, range = 1 -5-3-7 (1-9); dorsal fin height x = 24-8, range = 22-9-28- 1 (25-5); 
mental disc maximum length x = 5-4, range = 4-7-6-3 (5-5); mental disc maximum width x = 7-0, 
range = 5-7-7-7 (7-3); distance between snout and anus x = 71-4, range = 66- 6-75-0 (70-2); distance between 
snout and anal fin origin x = 74-3, range = 69-3-77-0 (72-2); distance between snout and pelvic fin origin 
x = 50-6, range = 47-9-52-6 (50-4); distance between snout and dorsal fin origin x = 45-2, range = 43-2-46-8 
(43-5). 

The body has a characteristic, streamlined shape (Fig. 6 and Krupp, 1983: fig. 30). From the 
pointed snout, the dorsal profile rises smoothly to the insertion of the dorsal fin. Behind the dorsal 
fin, the trunk diminishes in depth, terminating in a slender caudal peduncle almost exactly half as 
deep as long. In five specimens the abdomen had collapsed, so a reliable body depth measurement 
could not be taken. In ventral view, the upper lip is thick and has many small papillae. The shape of 




Fig. 6. Garra lautior. Holotype. Scale = 10 mm. 



66 



K. E. BANISTER 




Fig. 7. Details of the distribution of papillae on the frenum, lips and disc of (left) a paratype of Garra 
mamshuqua (74 mm SL ex 1976.4.7: 381-387), and (right) a paratype of Garra lautior (74 mm SL ex 
1976. 4.7: 399^04) Scale = 0-1 mm. 




Fig. 8. A scale from the row above the lateral line of Garra lautior. Scale = 0-5 mm. 



the disc and the disposition of the papillar beds are shown in Figs 2 & 7B. None of the specimens 
has any tubercles on the head although both mature males and females are present in the sample 
(see below). 

The small size range of the specimens available was insufficient to establish any marked instances 
of allometric growth. 

SQUAMATION. The scales are well developed and slightly lobate. A scale and its striations are shown 
in Fig. 8. Only two poorly defined growth rings were discernible. In the lateral line series there are 
32(f3), 33(f4), 34(f4) or 35(f2) scales. From the dorsal mid-line to the lateral line there are 3^(f9) or 
4|(f3) scales and from the lateral line to the pelvic fin base 3^(fl 3) scales. In front of the anal fin the 



SPECIES OF GARRA 



67 



ventral surface is scaleless. There are 12(fl3) scale rows around the least circumference of the 
caudal peduncle. 

VERTEBRAE. In the nine specimens radiographed there are 27(f2), 28(f3), 29(f3) or 30(fl ) vertebrae, 
excluding those comprising the Weberian mechanism. The abdominal vertebrae number 12(f3) or 
1 3(f6) (allowing for the difficulty in identifying the first caudal vertebra). The neural spines below 
the dorsal fin pterygiophores display the normal alignment and reduction in size of most Garra spp 
(Fig. 9) in contrast to the unique condition in Garra dunsirei (Fig. 4). 

In all the other species of Garra radiographed: viz G. mamshuqua Krupp, 1983, G. barreimiae 
Fowler & Steinitz, 1956, G. sahilia Krupp, 1983, G. tibanica Trewavas, 1941, and G. dunsirei, there 
are 4 interhaemal spine spaces corresponding to the anal fin pterygiophores (Fig. 10), but in G. 
lautior only three interhaemal spine spaces do so correspond (Fig. 1 1). No particular significance is 
attached to variations in the shape of the last anal fin pterygiophore. There are 1 4(f7) or 1 5(f2) pairs 
of ribs. 

GILL RAKERS. The gill rakers are small, hooked and number ll(fl), 12(fl), 13(f3), 14(fl), 15(f3), 
16(fl) and 17(fl) on the lower limb of the first gill arch. 

PHARYNGEAL BONES AND TEETH. The pharyngeal teeth number 2.4.5-5.4.2 (Fig. 12). The crowns 
have shallow spoon-edged depressions, the depression being most sharply edged in newly replaced 
teeth. 




Fig. 9. The vertebral column below the dorsal fin of Garra lautior to show the shape of the neural spines. 

Scale = 1 mm. 




Fig. 10. The anal fin pterygiophores of an unregistered BMNH specimen of Garra mamshuqua, 54 mm 
SL, to show their opposition to four interhaemal spaces. Scale = 1 mm. 



68 



K. E. BANISTER 




Fig. 11. The anal fin pterygiophores of a specimen of Garra lautior (unregistered) to show their 
opposition to only three interhaemal spaces. Scale = 1 mm. 




Fig. 12. Left pharyngeal bone of the holotype of Garra lautior. Scale = 0-5 mm. 



COLORATION. Alcohol preserved specimens are a uniform sandy brown, darker dorsally. The 
post-opercular spot is a deeper brown, but the 'Garra ' marks at the base of the dorsal fin membrane 
are not especially conspicuous. The fin membranes are clear. 

DISTRIBUTION. This species is known only from localities within the Wadi Hadramut drainage, 
Yemen. 

ETYMOLOGY. The trivial name is the comparative oflautus, the Latin for smart or neat and alludes 
to the neat, streamlined appearance of the fish. 

DIAGNOSIS. Garra lautior is sympatric only with Garra mamshuqua (see below). Although the two 
species are somewhat similar in body shape, Garra mamshuqua can be distinguished by the pres- 
ence of tubercles on the snout, the very conspicuous 'Garra ' marks on the dorsal fin and behind the 
operculum and the different disc shape (Fig. 2b & 7b). Very small specimens can be most easily 
separated on the greater intensity of the post-opercular spot in Garra mamshuqua. 



SPECIES OF GARRA 69 

TUBERCLES. Tubercles on the snout are often called nuptial or breeding tubercles (e.g. Wiley & 
Collette, 1970) or multicellular horny tubercles (Roberts, 1982). The latter author points out that 
they may occur in both sexes as well as being present before the onset of sexual maturity in Labeo 
species and the homalopterids. In Garra mamshuqua, the tubercles are present in both males and 
females at all stages of sexual maturity and first appear in specimens of 27 mm SL (e.g. in BMNH 
1967.4.7: 407-418). It seems probable therefore that in Garra mamshuqua the tubercles do not have 
a solely sexual or reproductive function. 

A hydrodynamic function was suggested by Reid (1978) for their occurrence in Labeo, since 
fishes from faster flowing waters had more and larger tubercles than those from quiet waters. The 
localities where the smooth Garra lautior and the tuberculate Garra mamshuqua are sympatric were 
described by the collector (original letter in the BMNH Fish Section archives) as 'a clear stream 
with stony shallows and deep holes' (Gheil Umar), and also an 'isolated muddy pot-hole below a 
dam' (Al-Ghurf). Such scanty and seemingly inconsistent information adds nothing to Reid's 
hydrodynamic hypothesis. 

Although the function of the tubercles is not known, it does seem in this case that their presence 
can be used as a sound diagnostic character to distinguish these two sympatric species. However, it 
is not suggested that the presence or absence of tubercles is diagnostic for other species. 



Discussion 

Krupp ( 1 983) also recorded both Garra tibanica and Garra sahilia from the Wadi Hadramut. Garra 
tibanica was included as a member of the Hadramut fauna solely on specimens collected by 
Scortecci at Bir el Manzil (1432'N, 4&5\'Ejide Krupp). Scortecci's Bir el Manzil is shown on 
the map in Balletto & Spano (1977), which is concerned with the Scortecci expedition and is 
approximately 1430'N, 4430'E or well to the west of the Hadramut. 

The Wadi Hadramut record of Garra sahilia was based on four fishes, two from Sayun (BMNH 
1980.4.24: 8,9) and two from nearby Shibam (BMNH 1980.4.24: 6,7). The latter specimens 
were identified by Krupp but not listed in his 1983 paper. The four fishes do not correspond to 
the description, especially in having much longer barbels and the anus closer to the anal fin than 
in Garra sahilia. However, their poor condition precludes confirmation of Krupp's specific 
determination. 

There are some difficulties in establishing which specimens of Garra sahilia are types. Krupp 
(1983: 601) lists 63 specimen (BMNH 1976.4.7:419,420-425; 1951.5.9: 12-65 and 1944.4.3: 1-10) 
as paratypes but used only 25 specimens in his description. Presumably, the 63 listed paratypes 
included the 24 actually described (although the largest specimen in his sample, 100- 5 mm SL 
BMNH 1940.2.15: 12-18 was used in the description but not designated a paratype). 

Although twice as many Garra lautior specimens were available to me than to Krupp, in 
most respects our descriptions are similar. However, there is substantial discrepancy in our scale 
counts around the caudal peduncle. In the 13 specimens used here (6 of which were those used by 
Krupp) I could count only 12 scales, whereas Krupp gives 14(fl) or 16(f5). A similar discrepancy 
occurs with the same count in Garra buettikeri Krupp, 1983, Krupp giving 18(f2) or 20(fl8) as the 
diagnostically high circumpeduncular scale count, whereas in the six BMNH specimens he used in 
his description I count only 16(f2) or 18(f4) scales. In Krupp's fig. 21 twenty scales would be too 
many, unless the squamation in the specimen illustrated was unusually asymmetrical. Although 
Krupp did not indicate how he made his counts I can imagine only one way of counting the number 
of scale rows around the least circumference of the caudal peduncle. It seems unwise, therefore, to 
attribute diagnostic significance to this particular meristic feature. No attempt is made in this paper 
to produce a key to the Garra species of the Arabian peninsula. Even a cursory glance at the means 
and ranges of any particular morphometric or meristic feature used here, in Krupp (1983), in 
Banister & Clarke (1977) and in many other papers shows that the similarity of means and the 
extensive overlaps in range usually precludes the use of such characters in a key. Even if the eight 
peninsular species were initially subdivided by drainage regions (giving groups of 3, 2, 2 and 1) a 
key based on morphometric and meristic characters would not infallibly separate the sympatric 



70 K. E. BANISTER 

species. The major diagnostic features are regrettably very difficult to quantify. At the moment the 
most useful characters are the overall body shape, the shape of the mental disc and the distribution 
of papillae thereon, and the colour pattern. Although details of the mental disc serve to distinguish 
species, based on the samples available, the variation in at least one species, Garra tibanica (Balletto 
& Spano, 1977: fig. 6) makes one wonder whether it will remain useful when more populations are 
discovered. 

Krupp 1983: 603-615 provided a useful list of all the specimens he examined. There are, 
however, some confusions in the BMNH register numbers in his list and these and some other 
errors are corrected below. 

Garra sahilia sahilia (p. 603) 

Sample No. 10 for Wadi Abd read Wadi Anad 4450'E, 1317'N 

No. 14 for 1976.4.7: 443^60 read 1976.4.7: 460 

No. 15 for 1910.1.28: 1-3 read 1870.1.28: 1-3 

Garra sahilia gharbia (p. 604) 

Sample No. 5 for 1976.4.7: 646-354 read 1976.4.7: 346-349 

Garra tibanica tibanica (p. 608) 

Sample No. 5 for 1976.4.7: 443^60 read 1976.4.7: 443^59 

No. 1 1 for 1952.5.7: 13-18 read 1952.5.7: 13-17 

No. 17 for 1976.4.7: 346-356 read 1976.4.7: 350-354 

Garra incertae sedis (p. 615) 

Sample No. 1 for 1976.4.7: 374-377 read 1976.4.7: 377 

No. 3 for 1976.4.7: 380-406 read 1976.4.7: 380-396 



Acknowledgements 

I particularly wish to thank the collectors for their enthusiasm in seeking for fish life in such an inhospitable 
environment. Without the efforts of such people, our knowledge of fish distribution and habitats would 
increase more slowly. My colleagues, Dr P. J. P. Whitehead and Mr A. C. Wheeler offered constructive 
comments on the manuscript. My thanks go to them and to Gordon Howes for illustrating the new species 
and to Joan Ellis for typing the paper. 



Bibliography 

Balletto, E. & Spano, S. 1977. Ciprinidi del genere Garra Hamilton 1822, raccolti nello Yemen dal Prof. 

Guiseppe Scortecci. Annali del Museo Civico di Storia Naturale (di Genova) Giacomo Doria, Genova 81: 

246-287. 
Banister, K. E. 1984. A subterranean population of Garra barreimiae (Teleostei: Cyprinidae) from Oman, 

with comments on the concept of regressive evolution. Journal of Natural History 18: 927-938. 
Banister, K. E. & Clarke, M. A. 1977. The freshwater fishes of the Arabian peninsula. The scientific results of 

the Oman Flora and Fauna survey 1975. Journal of Oman Studies 1977: 1 1 1-154. 
Krupp, F. 1983. Fishes of Saudi Arabia and adjacent regions of the Arabian Peninsula. Fauna of Saudi Arabia 

5: 568-636. 
Reid, G. McG. 1978. A systematic study oflabeine cyprinid fishes with particular reference to the comparative 

morphology and morphometrics of African Labeo species. Ph.D. thesis, University of London, 770 pp. 
Roberts, T. R. 1982. Unculi (horny projections arising from single cells), an adaptive feature of the epidermis 

of ostariophysan fishes. Zoologica Scripta 11: 55-76. 
Wiley, M. L. & Collette, B. B. 1970. Breeding tubercles and contact organs in fishes: their occurrence, 

structure and significance. Bulletin of the American Museum of Natural History 143: 143-216. 

Manuscript accepted for publication 6 November 1985 



British Museum (Natural History) 

The birds of Mount Nimba, Liberia 



Peter R. Colston & Kai Curry-Lindahl, with a section on the biogeographic 
context by Malcolm Coe. 



For evolution and speciation of animals Mount Nimba in Liberia, Guinea and the Ivory Coast is 
a key area in Africa representing for biologists what the Abu Simbel site in Egypt signified for 
archaeologists. No less than about 200 species of animals are endemic to Mount Nimba. Yet, this 
mountain massif, entirely located within the rain-forest biome, is rapidly being destroyed by 
human exploitation. 

This book is the first major work on the birds of Mount Nimba and surrounding lowland 
rain-forests. During 20 years (1962-1982) of research at the Nimba Research Laboratory in 
Grassfield (Liberia), located at the foot of Mount Nimba, scientists from three continents have 
studied the birds. In this way Mount Nimba has become the ornithologically most thoroughly 
explored lowland rain-forest area of Africa. 

The book offers a comprehensive synthesis of information on the avifauna of Mount Nimba 
and its ecological setting. During the 20 years period of biological investigations at Nimba this in 
1962 intact area was gradually opened up by man with far-reaching environmental consequences 
for the rain-forest habitats and profound effects on the birds. Therefore, the book provides not 
only a source of reference material on the systematics, physiology, ecology and biology of the 
birds of Mount Nimba and the African rain-forest, but also data on biogeography in the African 
context as well as on conservation problems. Also behaviour and migration are discussed. At 
Nimba a number of migrants from Europe and/or Asia meet Afrotropical migratory and 
sedentary birds. 

Professor Kai Curry-Lindahl has served as Chairman of the Nimba Research Laboratory and 
Committee since its inception in 1962. Peter Colston is from the Subdepartment of Ornithology, 
British Museum (Natural History), Tring, and Malcolm Coe is from the Animal Ecology 
Research Group, Department of Zoology, Oxford. 

1986, 129pp. Hardback. 565 00982 6 17.50. 



Titles to be published in Volume 52 

Miscellanea 

A revision of the Suctoria (Ciliophora, Kinetofragminophora) 5. The Paracineta 
and Corynophom problem. By Colin R. Curds 

Notes on spiders of the family Salticidae 1. The genera Spartaeus, Mintonia and 
Taraxella. By F. R. Wanless 

Mites of the genus Holoparasitus Oudemans, 1936 (Mesostigmata: Parasitidae) in 
the British Isles. By K. H. Hyatt 

The phylogenetic position of the Yugoslavian cyprinid fish genus Aulopyge Heckel, 
1841, with an appraisal of the genus Barbm Cuvier & Cloquet, 1816 and the 
subfamily Cyprininae. By Gordon J. Howes 

Revision of the genera Acineria, Trimyema, and Trochiliopsis (Protozoa, 
Ciliophora). By H. Augustin, W. Foissner & H. Adam 

The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) with a 
systematic review, a synopsis of Pipistrellus and Eptesicus, and the descriptions of 
a new genus and subgenus. By J. E. Hill & D. L. Harrison 

Notes on some species of the genus Amathia (Bryozoa, Ctenostomata). By P. J. 

Chimonides 



Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 



A revision of the Suctoria (Ciliophora, 
Kinetofragminophora) 5. The Paracineta 
and Corynophrya problem 

Colin R. Curds 



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Cromwell Road 
London SW7 5BD Issued 27 February 1987 



PRESENTED 



II f 
A revision of the Suctoria (Ciliophor a, f L 1 2 7 F E B 1987 

Kinetofragminophora) 5. The Paradneta and 
Corynophrya problem 






Colin R. Curds 

Zoology Department, British Museum (Natural History), Cromwell Road, London 




Contents 



Synopsis 
Introduction . 
Genus Actinocyathula 
Genus Corynophrya 
Genus Pelagacineta 
Genus Paradneta . 
Genus Loricophrya . 
Genus Anthacineta . 
Genus Flectacineta . 
References 
Index to species 



71 

71 

72 

76 

82 

86 

91 

98 

101 

104 

106 



Synopsis 



The continual drift in the diagnosis of the unrelated genera Paradneta and Corynophrya causes considerable 
taxonomic problems and confusion at several levels in classification. The transfer of Paradneta crenata and 
Paradneta homari into the genus Actinocyathula has allowed the present review to be based, as far as possible, 
on the original diagnoses of the genera. In addition to those mentioned above, the species of four other genera, 
Pelagacineta, Loricophrya, Anthacineta and Flectacineta are reviewed since some have been previously 
associated in some way with the Paradneta-Corynophrya problem in the past. 

A new diagnosis for each genus is given with a key to its constituent species and where appropriate a 
genotype has been designated to encourage taxonomic stability. All species are described and figured. 



Introduction 

There is still considerable confusion and disagreement on the generic diagnoses of Paradneta 
Collin, 1911 and Corynophrya Kahl, 1934. The purpose of this publication is to review the species 
involved, to amend previous diagnoses and to assign type species to the genera in an attempt to 
establish taxonomic stability. The genus Paradneta was erected in order to take account of those 
loricate suctoria with an apical group of tentacles that reproduced by external budding and that 
were longitudinally symmetrical. In his original generic description, Collin (1911) included the 
three species Paradneta crenata (Fraipont, 1878), P. homari (Sand, 1899) and P.patula (Claparede 
& Lachmann, 1861) but failed to designate the type species. In his later taxonomic revision, 
Collin (1912) transferred several more species into the genus including Paradneta limbata 
(Maupas, 1881), P. vorticelloides (Fraipont, 1878), P. jorisi (Sand, 1895), P.parva (Sand, 1899), 
P. multitentaculata (Sand, 1895), P. livadiana (Mereschkowsky, 1881), P. elegans (Imhoff, 1883) 
and P. bifaria (Stokes, 1887). Collin (1911, 1912) stressed that although external budding was a 
prime feature of the genus both Paradneta crenata and P. homari in fact reproduced by semi- 
external budding (the semi-invaginative budding of Batisse, 1975). At the time this method was 
thought to be only a slight variation on the external budding theme and of little significance. 



Bull. Br. Mas. not. Hist. (Zool.) 52(2): 71-106 



Issued 27 February 1987 



71 



72 C. R. CURDS 

Modern workers however consider the different modes of budding to be of great taxonomic 
importance and that there is a distinct difference between semi-invaginative and external budding. 
Nevertheless, the two species remained in their original genus until Batisse (1975) suggested their 
transfer into the genus Corynophrya Kahl, 1934 which had been originally erected for a hetero- 
genous assemblage of aloricate suctoria reproducing by internal budding. Although the suggestion 
by Batisse (1975) may appear strange, since the two species in question are loricate and reproduce 
differently, it should be pointed out that the generic diagnosis of Corynophrya has drifted consider- 
ably since that originally outlined by Kahl (1934). However, Batisse (1975) had not taken into 
account that Paracineta crenata can be regarded to be congeneric with Actinocyathus cidaris Kent, 
1882 and would be more neatly transferred into the latter older genus. Jankowski (1981) is also 
apparently of a similar opinion since he suggested that the name Actinocyathus might replace that 
of Paracineta. The name Actinocyathus was shown by Corliss (1960) to be preoccupied and he 
suggested the replacement name Actinocyathula Corliss, 1960. 

Kahl (1934) erected the genus Corynophrya to include the mostly marine assemblage of suctoria 
which Collin (1912) had gathered together in his third group within the genus Discophrya. The 
major diagnostic features were that they reproduced by internal budding, did not possess a lorica, 
were rounded in cross-section, had one type of tentacle that was restricted to the apical surface and 
had a rounded, compact nucleus. According to Kahl (1934) the following species held these 
features in common, Corynophrya marina (Andrusov, 1886), C. conipes (Mereschkowsky, 1879), 
C. macropus (Meunier, 1910), C. lyngbyi (Ehrenberg, 1833), C. francottei (Sand, 1895), 
C. campanula (Schroder, 1907), C. interrupta (Shroder, 1907) and C. stueri (Schroder, 1911). 
Kahl agreed with Collin (1912) and placed the genus in the family Discophryidae where it remained 
until Batisse (1975) transferred it into the Thecacinetidae. More recently Jankowski (1978) has 
transferred three of the species, which clearly have elongate to branched macronuclei and multiple 
endogenous buds, into the new genus Pelagacineta Jankowski, 1978. 

Genus ACTINOCYATHULA Corliss, 1960 

Actinocyathus Kent, 1882 
Corynophrya sensu Batisse, 1975 
Paracineta sensu Jankowski, 1978 
Faltacineta Jankowski, 1982 

The genus Actinocyathus was erected by Kent (1882) for those resembling Ephelota in general 
form but borne upon a stalked lorica. Kent's (1882) diagnosis also stated that the tentacles were 
retractile but not capitate. However, Kent further stated in his description of the type species 
Actinocyathus cidaris Kent, 1882 that he only saw the tentacles in the contracted state which 
leaves the absence of capitate tentacles open to considerable doubt. There seems to be little doubt 
that the organism depicted by Dons (1922) which he calls Paracineta crenata (Fraipont) forma 
pachyteca Collin (Dons mispelling ofpachytheca) is congeneric with Actinocyathus and conspecific 
with Acineta crenata Fraipont, 1878. In view of this the two species Paracineta crenata (Fraipont, 
1878) and P. homari (Sand, 1899) which both reproduce by semi-invaginative budding are trans- 
ferred to Actinocyathula Corliss, 1960. Jankowski (1982) erected the genus Faltacineta Jankowski, 
1982 for the two marine epizoic species Paracineta pleuromammae Steuer, 1928 and Paracineta 
gaetani Sewell, 1951. However, the former species P. pleuromammae is clearly depicted showing 
semi-invaginative budding and for this reason the two are transferred to Actinocyathula for the 
first time. 

Diagnosis of Actinocyathula 

Marine suctorians whose ovoid-shaped body is restricted to the anterior half of the lorica. 
Lorica cup-shaped, never laterally compressed, borne upon a stalk and attached to marine 
invertebrates such as Crustacea, hydroid colonies and calcareous sponges. Tentacles in a single 
group that is restricted to the apical region of the body. Actinophores absent. Reproduction by 
semi-invaginative budding. 



PARACINETA AND CORYNOPHRYA PROBLEM 

Key to the species of Actinocyathula 

1 Stalk equal to or less than lorica length, epizoic on Crustacea 

Stalk greater than lorica length, epizoic on invertebrates other than Crustacea 

2 Lorica smooth 

Lorica striated transversely . . . . . ... 

3 Posterior region of lorica broadly rounded 

Posterior of lorica distinctly narrow 

4 Lorica elongate, stalk usually less than half lorica length .... 
Lorica width and stalk length approximately equal to lorica length 



73 



3 

.. . .2 

. A. cidaris 

. A. crenata 

. A. ho mar i 

. 4 

A . pic lit- o mammae 
. A.gaetani 



Species descriptions 

Actinocyathula cidaris Corliss, 1960 
Actinocyathus cidaris Kent, 1882 

DESCRIPTION (Fig. 1). This the type species is a small (40 ^m long), marine, loricate suctorian. The 
ovoid body has a flattened base and protrudes from the apical region of the lorica. Tentacles 
retractile, radiating from the anterior surface of body . Lorica surface smooth, triangular in outline, 
rounded in cross-section. Apical edge of lorica bends inwards to form a thin cup-like platform in 
which the zooid is located. Lorica mounted on slender but rigid stalk that is 3-4 times the lorica 
length. Epizooic on the calcareous sponge Grantia compressa. Contractile vacuole may be single or 
double. Nuclear and reproductive features not described. 




Fig. 1 Actinocyathula cidaris after Kent, 1 882 (called Actinocyathus cidaris). 

Actinocyathula crenata n. comb. 

Acineta crenata Fraipont, 1878 

Acineta saifulae Mereschkowsky, 1877 

Paracineta crenata Collin, 1911 

Paracineta crenata var. pachytheca Collin, 1912 

Paracineta crenata forma pachyteca Dons, 1922 

Corynophrya crenata Batisse, 1975 

Miracineta saifulae Jankowski, 1981 



74 



C. R. CURDS 







Fig. 2 Actinocyathula crenata: (a-c) after Collin, 1912 (called Paracineta crenata); (d,e) after 
Mereschkowsky, 1877 (called Acineta saifulae); (f) after Collin, 1912 (called Paracineta crenata); 
(g) after Fraipont, 1878 (called Acineta crenata); (h) after Dons, 1922 (called Paracineta crenata var. 
pachytheca); (i) after Wailes, 1928 (called Paracineta crenata var. pachytheca). 



DESCRIPTION (Fig. 2). Medium (75 |im long), marine, loricate suctorian. The ovoid body protrudes 
from the apical region of the lorica. Capitate tentacles sometimes retractile, radiating from the 
anterior surface of body. Lorica surface crenulated with three to many transverse striations, 
triangular to elongate in outline, rounded in cross-section. There is a thin cup-like platform in 
which the zooid is located. Lorica mounted on slender stalk that is 3-4 times the lorica length. 
Epizooic on a variety of marine invertebrates including the hydroids Clytia volubilis, Leptoscyphus 
grigoriewi and Perigonimus repens and the polychaete Aphrodite aculeata. Single contractile 
vacuole located laterally. Spherical macronucleus centrally positioned. Reproduction by semi- 
invaginative budding. Swarmer not described. 



PARACINETA AND CORYNOPHRYA PROBLEM 



75 





Fig. 3 Actinocyathula gataeni: (a-d) various growth stages; (e-g) adults; all after Sewell, 1951 (called 

Paracineta gataeni). 



A ctinocyathula gataeni (Sewell, 1951) n. comb. 

Paracineta gataeni Sewell, 195 1 
Faltacineta gataeni Jankowski, 1 982 

DESCRIPTION (Fig. 3). Small (30-55 um diameter), marine, loricate suctorian. The ovoid body 
protrudes from the apical region of the lorica. Tentacles radiate out from the anterior body surface. 
Lorica surface usually smooth but sometimes with transverse wrinkles, triangular in outline, 
rounded in cross-section. Lorica mounted on a robust rigid stalk that is usually less than the lorica 
length. Lorica sometimes mounted eccentrically on stalk. Epizooic on the copepods Gaetanus 
antarcticus Wolfendon and G. curvicornis Sars. Macronucleus spherical. Reproduction and 
swarmer not described. 

Actinocyathula homari n. comb. 

Acineta homari Sand, 1899 
Paracineta homari Collin, 1911 
Corynophrya homari Batisse, 1975 

DESCRIPTION (Fig. 4). Small (25-40 urn long), marine, loricate suctorian. The ovoid body 
protrudes from the apical region of the lorica. Tentacles retractile, radiating out from the anterior 
body surface. Lorica surface smooth, triangular to bell-shaped in outline, rounded in cross- 
section. Lorica mounted on a robust rigid stalk that rarely exceeds the lorica length. Lorica 
sometimes mounted eccentrically on stalk. Epizooic on a variety of decapod Crustacea. Single 
contractile vacuole located centrally or laterally. Macronucleus spherical, located at posterior of 
body. Reproduction by semi-invaginative budding. Swarmer not described. 



76 



C. R. CURDS 





Fig. 4 Actinocyathula homari: (a-d) after Collin, 1912 (called Paracineta homari); (e,f) after Sand, 1 899 

(called Acineta homari}. 



Actinocyathula pleuromammae (Steuer, 1928) n. comb. 

Paracineta pleuromammae Steuer, 1928 
Faltacineta pleuromammae Jankowski, 1982 

DESCRIPTION (Fig. 5). Medium (60-1 15 jam long), marine, loricate suctorian. The ovoid body 
protrudes from the apical region of the lorica. Tentacles radiate out from the anterior body surface. 
Lorica surface with irregular transverse striations, elongated cone, rounded in cross-section. 
Lorica mounted on a robust rigid stalk that is less than half the lorica length. Epizoic on the 
copepods Pleuromamma abdominalis and P. xiphias. Single contractile vacuole located laterally. 
Macronucleus spherical, located centrally. Reproduction by semi-invaginative budding. Swarmer 
ovoid with many transverse ciliary rows. 

Genus CORYNOPHRYA Kahl, 1934 
Pelagacineta Jankowski, 1 978 pro pane 

The genus was orginally erected by Kahl (1934) to include a heterogenous collection of mainly 
marine species. He stated that the major features distinguishing it from other genera included 
internal budding, a single apical group of tentacles and a rounded, compact macronucleus. Kahl 
(1934) included eight species in his genus but three have recently been transferred to the new genus 
Pelagacineta by Jankowski (1978). Kahl (1934) followed the original higher classification system 
of Collin (1912) and placed the genus in the family Discophryidae where it remained until Batisse 
(1975) transferred it into the Thecacinetidae which demands reproduction by semi-invaginative 
budding. The latter step was taken because Batisse (1975) had included Actinocyathula 
(Paracineta) crenata and A. homari in the genus. In fact the mode of budding has only been 
described for one of the five remaining species, where in Corynophrya lyngbyi it is endogenous. 



PARACINETA AND CORYNOPHRYA PROBLEM 



77 





10 



Fig. 5 Actinocyathula pleuromammae: (a-c) after Steuer, 1928 (called Paracineta pleuromammae). 



However, Jankowski (1981) was recently of the opinion that genera in his family Corynophryidae 
reproduce exogenously although he gave no practical evidence for that conclusion. Of those 
which Kahl (1934) originally included in the genus only four, Corynophrya macropus, C. conipes, 
C. lyngbyi and C. francottei remain in the present review. The anterior notch in the body of 
Corynophrya marina has been interpreted to indicate invaginative budding and will be transferred 
to an appropriate genus in a later publication. One other species, Ephelota columbiae Wailes, 1943 
is included in the genus for the first time since it bears only one type of tentacle whereas there are 
two types in Ephelota. The five species that are included have several features in common, they all 
have a compact rounded macronucleus, a single apical group of tentacles that are both retractile, 
prehensile and suctorial and in most there is a conical stalk that clearly narrows towards its base. 
The species most completely described is Corynophrya lyngbyi and this is designated to be the type 
species in an attempt to establish taxonomic stability. 

Diagnosis of Corynophrya 

Mainly marine, aloricate suctorians whose body shape is spherical to ovoid, rounded in cross 
section. Borne upon a stalk which is commonly stout near to the zooid narrowing markedly 
towards its base. Usually epizooic on hydroids, Crustacea and polychaetes but also noted on 
marine algae. Tentacles prehensile and retractile in a single group that is restricted to the 
apical region on the body. Actinophores absent. Macronucleus usually spherical. Reproduction by 
endogenous budding. 



Key to the species of Corynophrya 

1 Stalk long, at least 3 times length of body 
Stalk short, up to twice length of body 



78 C. R. CURDS 

2 Freshwater, tentacles wide at base, narrowing towards capitate ends . 
Marine, sides of tentacles parallel, do not narrow towards capitate ends 

3 Body spherical and regular ........ 

Body ovoid, uneven with folds 

4 Stalk striated transversely 

Stalk striated longitudinally or without striations .... 

5 Macronucleus spherical 

Macronucleus in shape of horseshoe 

6 Stalk markedly wider near zooid, narrowing towards base . 
Sides of stalk parallel, stalk does not narrow towards base . 



. C. tumida 
3 

C. columbine 

C. symbiotica 

. C. conipes 

5 

. 6 

. C. lyngbyi 

C. macropus 

C.francottei 



Species descriptions 

Corynophrya lyngbyi (Ehrenberg, 1833)Kahl, 1934 

Acineta lyngbyi Ehrenberg, 1833 

Podophrya lyngbyei Claparede & Lachmann, 1859 non Robin, 1879 

Tokophrya lyngbyei Biitschli, 1889 

Discophrya lyngbyei Collin, 1912 

DESCRIPTION (Fig. 6). This the type species is a small to medium (40-80 urn), marine, aloricate 
suctorian. The ovoid body is oval in section and slightly wider anteriorly. The retractile, capitate 
tentacles located on the anterior body surface. Stalk long (1 20-400 um), at least four times 




Fig. 6 Corynophrya lyngbyi after Fraipont, 1 878 (called Podophrya lyngbyi). 



PARACINETA AND COR YNOPHR YA PROBLEM 79 




10 



Fig. 7 Corynophrya columbiae after Wailes, 1943 (called Ephelota columbiae}. 



the body length. Stalk wider near zooid than at its base. Attached to hydroid colonies such as 
Sertularia and Clytia as well as marine algae. There are one or two contractile vacuoles. 
Macronucleus spherical in the young adult but this elongates into a horse-shoe shape at maturity. 
Reproduction by endogenous budding which may be multiple. Swarmer not described. 

NOTE. The specific epithet has been consistently mispelt by several authors over many years. 
Ehrenberg's (1833) original spelling was lyngbyi but later (1838) in his atlas the name appears as 
lyngbyei and it was this spelling that was used by several later authorities. 

Corynophrya columbiae n. comb. 
Ephelota columbiae Wailes, 1943 

DESCRIPTION (Fig. 7). This is a small (30-60 um), marine, aloricate suctorian. The spherical to 
ovoid body is round in section. The retractile, capitate tentacles located on the anterior half of 
body surface. Stalk usually short (50-200 um), and usually less than three times the body length. 
Stalk wide near zooid narrowing towards the base. Attached to Crustacea in large numbers. 
Macronucleus spherical, centrally located. Reproduction not described. 

Corynophrya conipes (Mereschkowsky, 1877)Kahl, 1934 

Acineta conipes Mereschkowsky, 1877 
Podophrya conipes Mereschkowsky, 1879 
Tokophrya conipes Biitschli, 1889 

DESCRIPTION (Fig. 8). This is a large (100-190 um), marine, aloricate "suctorian. The ovoid to 
pyriform body is oval in section and widens anteriorly. The retractile, capitate tentacles located 
mainly on the anterior body surface. Stalk long (800-1 500 urn), usually 8-10 times the body length. 
Stalk distinctly wider near zooid than at its base, finely striated transversely and usually with two 
distinct annuli situated about a third of the way down the stalk. Attached to marine algae such as 
Ptilota and Ceramium. Single anterior contractile vacuole. Macronucleus spherical, located 
centrally or subcentrally. Reproduction and swarmer not described. 

Corynophryafrancottei(Sand, 1895)Kahl, 1934 

Tokophrya francottei Sand, 1895 
Discophryafrancottei Collin, 1912 

DESCRIPTION (Fig. 9). This is a small (50-60 um), marine, aloricate suctorian. The retractile, 
capitate tentacles are located on the anterior surface of the spheroidal body. Stalk long 
(1 00-230 um), at least three times the body length, retaining a constant diameter along its 
entire length. Attached to hydroid colonies such as Sertularia and Ceramium. There is a single 



80 



C. R. CURDS 




Fig. 8 Corynophrya conipes: (a) after Mereschkowsky, 1879 (called Podophrya conipes); (b) after 

Meunier, 1910 (called Podophrya conipes). 




10 



Fig. 9 Corynophrya francottei after Sand, 1895 (called Tokophryafrancottei). 



PAR A CINETA AND COR YNOPHR YA PROBLEM 8 1 




Fig. 10 Corynophrya macropus after Meunier, 1910 (called Podophyra macropus). 

marginal contractile vacuole. Macronucleus oval to spherical, located centrally or subcentrally. 
Reproduction and swarmers not described. 

Corynophrya macropus (Meunier, 1910)Kahl, 1934 
Podophrya macropus Meunier, 1910 

DESCRIPTION (Fig. 10). This is an incompletely defined species whose size has not been recorded, 
marine, aloricate. The body is spherical in shape and carries retractile, capitate tentacles on its 
anterior surface. Stalk long, at least three times the body length. Stalk, which is wider near the 
zooid than at its base, is distinctly striated, longitudinally along its entire length. Macronucleus 
spherical, located centrally. Reproduction and swarmer not described. 

Corynophrya symbiotica Jankowski, 1981 

DESCRIPTION (Fig. 1 1). This is a medium (80-105 urn), marine, aloricate suctorian. The ovoid body 
has rather bumpy irregular appearance with some longitudinal folds. The retractile tentacles 
occupy the entire domed anterior body surface. Stalk comparatively short (up to 90 jam), about 
same as the body length. Stalk slightly wider near zooid than at its base. Attached to arctic 
polychaete worms belonging to the family Aphroditidae. There is a single anterior contractile 
vacuole. Macronucleus spherical, located centrally. Reproduction and swarmer not described. 

Corynophrya tumida (Gajewskaja, 1933) Matthes, 1954 
Discophrya tumida Gajewskaja, 1933 

DESCRIPTION (Fig. 12). This is a small (50 um), freshwater, aloricate suctorian. The ovoid body is 
round in section and slightly wider posteriorly. The retractile, capitate tentacles are rather wider 
at the base and occupy the anterior half of the body surface. Stalk short (60-70 urn), only just 
longer than the body. Stalk wider near zooid than at its base and distinctly striated transversely 
at infrequent intervals along its length. The stalk is also irregularly striated longitudinally. 



82 



C. R. CURDS 




10 



Fig. 1 1 Corynophrya symbiotica after Jankowski, 1 98 1 , 




Fig. 12 Corynophrya tumida after Gajewskaja, 1933 (called Discophrya tumidd). 



Attached to gammarid Crustacea in Lake Baikal. There is a single anterior contractile vacuole. 
Macronucleus spherical, located centrally. Reproduction and swarmer not described. 

Genus PELAGACINETA Jankowski, 1978 

Schroder (1907) first described the two marine species Tokophrya interrupta and T. campanula 
which resembled Ephelota in some respects and Podocyathus in others. They resembled Ephelota in 
their multiple endogenous method of budding but Ephelota is without a thecostyle and has two 
different types of tentacles. Similarly they resembled Podocyathus in their overall structure but 
reproduced differently from that genus. Schroder (191 1) later added a further species T. steueri to 
the group but still placed it in Tokophrya a genus typified by the absence of a lorica. Collin (1912) 
was the first to transfer the three species out of Tokophrya and he grouped them with several other 
misfits into his third section of the genus Discophrya. Kahl (1934) later erected the new genus 
Corynophrya for Collin's third section where they remained until the genus Pelagacineta was 



PAR A CINETA AND COR YNOPHR YA PROBLEM 8 3 

defined by Jankowski (1978) for those species 'like Podocyathus but with multiple endogenous 
budding'. Jankowski (1978) designated P. interrupta (Schroder, 1907) to be the type species and 
included P. campanula (Schroder, 1907) in the new genus. In the current revision the diagnosis is 
elaborated for the sake of clarity and some other species are transferred to the genus for the first 
time. 

Diagnosis of Pelagadneta 

Marine suctoria with lorica-like thecostyle. Body shape ovoid, discoidal or pyriform, rounded in 
cross section, actinophores absent. Stalk widens anteriorly to form lorica-like thecostyle. Single 
type of retractile tentacle present, arranged in one or two anterior groups. Attached to copepods or 
marine algae. Macronucleus typically elongate and often branched. Reproduction by multiple 
endogenous budding. Swarmers ovoid partially ciliated with several longitudinal kinetics. 

Key to the species of Pelagadneta 

1 Tentacles in single anterior group P. campanula 

Tentacles in two anterior groups 2 

2 Only 2 tentacles present, attached to algae P. dibdalteria 

Many tentacles present, attached to copepods 3 

3 Macronucleus elongate but not branched, body ovoid but not discoidal . . . P. euchaetae 
Macronucleus elongate and branched, body sometimes discoidal .... P. interrupta 



Species descriptions 
Pelagadneta interrupta (Schroder, 1907) Jankowski, 1978 

Tokophrya interrupta Schroder, 1907 
Discophrya interrupta Collin, 1912 
Corynophrya interrupta Kahl, 1934 

DESCRIPTION (Fig. 1 3). This the type species is a medium (100-140 urn long), marine suctorian with 
thecostyle. The ovoid body may be dorso-ventrally compressed and discoidal in shape lying at the 
top of a thecostyle that widens considerably to form a lorica-like anterior region. Stalk region 
hollow, 2-3 times the length of the lorica part of the thecostyle, terminating in a longitudinally 
striated basal disc. Many retractile, capitate tentacles located anteriorly arranged in two fascicles. 
Attached to marine copepods such as Euchaeta and Metridia reported from antarctic waters. 
Shape of macronucleus variable, always elongate and frequently branched. Reproduction by 
multiple endogenous budding producing oval swarmers partially ciliated with many kinetics on 
part of the ventral body surface. 

Pelagadneta campanula (Schroder, 1907) Jankowski, 1978 

Tokophrya campanula Schroder, 1907 
Tokophrya steueri Schroder, 1911 
Discophrya campanula Collin, 1912 
Discophrya steueri Collin, 1912 
Corynophrya campanula Kahl, 1934 
Corynophrya steueri Kahl, 1934 

DESCRIPTION (Fig. 14). This is a medium (100-1 50 um long), marine suctorian with thecostyle. The 
ovoid body may be dorso-ventrally compressed and discoidal in shape lying at the top of a 
thecostyle that widens considerably to form a cupped lorica-like anterior region. Stalk region 
hollow, 1-3 times the length of the lorica part of the thecostyle, terminating in a longitudinally 
striated basal disc. Many retractile, capitate tentacles located anteriorly arranged in a single 
fascicle sometimes surrounded by an outer ring of short tentacles. Attached to marine copepods 
such as Euchaeta and Metridia reported from antarctic waters. Shape of macronucleus variable but 



84 



C. R. CURDS 





Fig. 13 Pelagacineta interrupta: (a,b) after Schroder, 1907 (called Tokophrya interruptd). 



always elongate and highly branched. Reproduction by multiple endogenous budding producing 
oval swarmers partially ciliated with many kineties on part of the ventral body surface. 

Pelagacineta dibdalteria (Parona, 1881), n. comb. 
Acineta dibdalteria Parona, 1881 

DESCRIPTION (Fig. 1 5). This is a small (50-60 um long), marine suctorian with thecostyle. The body 
is pyriform in outline, rounded in cross section and lies at the top of a thecostyle that widens 
considerably to form a cupped lorica-like anterior region. Stalk region hollow, equal to or slightly 
less than the length of the lorica part of the thecostyle. There are only two capitate, prehensile 
mobile tentacles, one located anteriorly on either side of the body. Attached to marine algae. 
Contractile vacuole positioned centrally. Macronucleus elongate sausage-shaped. Reproduction 
and swarmers not described. 



Pelagacineta euchaetae (Sewell, 1951) n. comb. 
Acineta euchaetae Sewell, 1951 

DESCRIPTION (Fig. 16). This is a medium (80-90 um diameter), marine suctorian with thecostyle. 
The ovoid body lies at the top of a thecostyle that widens considerably to form a lorica-like anterior 
region. Young forms without lorica portion of the thecostyle. Stalk region hollow, usually shorter 
than length of the lorica part of the thecostyle, terminating in a longitudinally striated basal disc. 
Many retractile, capitate tentacles located anteriorly arranged in two fascicles. Attached to the 



PARACINETA AND CORYNOPHRYA PROBLEM 



85 




Fig. 14 Pelagacineta campanula: (a-c) after Schroder, 1907 (called Tokophrya campanula); (d,e) adult 
and swarmer, after Schroder, 1911 (called Tokophrya steueri). 




10 



Fig. 15 Pelagacineta dibdalteria after Parona, 1881 (called Acineta dibdalteria). 



86 



C. R. CURDS 




Fig. 16 Pelagacineta euchaetae: various growth stages, after Sewell, 1951 (called Acineta euchaetae). 



marine copepod Euchaeta reported from antarctic waters. Shape of macronucleus variable, always 
elongate and curved. Reproduction by endogenous budding producing oval swarmers. 

Genus PARACINEIA Collin, 191 1 

Luxophrya Jankowski, 1978 
Proluxophrya Jankowski, 1978 
Stemacineta Jankowski, 1978 

The genus Paracineta Collin, 1911 was erected in order to provide for those loricate suctoria 
with an apical group of tentacles that reproduced by external budding and were longitudinally 
symmetrical. The inclusion of Paracineta crenata, and P. hotnari which reproduce by semi- 
invaginative budding has already been dealt with above, but even after their removal, the species 
included by Collin (1912) in the genus Paracineta form a heterogenous group. Several other 
transfers have been suggested and are dealt with in other parts of this paper. After the removal 
of these from the genus the following four species remain from Collin's (1912) list, Paracineta 
jorisi (Sand, 1895), P. limbata (Maupas, 1881), P. patula (Claparede & Lachmann, 1861) and 
P. vorticelloides (Fraipont, 1878). Since that time, one other valid species has been added. One of 
the remaining major problems is the lack of a type species that will give some stability to the genus 
and enable a modern diagnosis to be proposed. This omission is rectified here by designating 
Paracineta patula (Claparede & Lachmann, 1861) Collin, 1911 as type species for the genus. 
This species is well described and includes good illustrated accounts of the budding and general 



87 

morphology. Furthermore it is the only surviving species of the three originally placed in the genus 
by Coffin (1911). 

Diagnosis of Paracineta 

Marine suctorians whose body shape is spherical to ovoid, rounded in transverse section. Long 
thecostyle with a semi-lorica that is variable in size. Semi-lorica may be sufficient to enclose half the 
zooid's volume or be reduced sufficiently for the body to be perched on top of a small cone-like 
widening at the top of the stem. Capitate tentacles usually restricted to apical body face but may 
radiate out from other areas when the semi-lorica is very small. Reproduction by exogenous 
budding, swarmers covered in many transverse ciliary rows. 

Key to the species of Paracineta 

1 Zooid perched on top of very small semi-lorica 2 

Approximately half of zooid enclosed within semi-lorica 4 

2 Tentacles emerge from all over zooid 3 

Tentacles restricted to apical surface P.jorisi 

3 Zooid with thick gelatinous outer covering P. limbata 

Zooid without gelatinous outer covering P. vorticelloides 

4 Stem of thecostyle with narrow flexible portion near junction with zooid .... P.patula 
Stem of thecostyle not narrowed, not flexible 5 

5 Thecostyle striated transversely regularly along entire length P. moebiusi 

Thecostyle smooth, unstriated 6 

6 Semi-lorica with border-like rim P.jorisi 

Semi-lorica without border-like rim 7 

7 Small, (semi-lorica 1 5-25 urn long), epizoic on polychaetes P. irregularis 

Medium, (semi-lorica 30-80 urn long), epizoic on hydroids and marine algae . . . P.patula 



Species descriptions 

Paracineta pat ula (Claparede & Lachmann, 1861) Collin, 1911 

Acineta patula Claparede & Lachmann, 1861 
Acineta divisa Fraipont, 1878 
Paracineta divisa Kahl, 1934 
Stemacineta patula Jankowski, 1978 

DESCRIPTION (Fig. 17). This the type species is a small (50-60 um long), marine suctorian with a 
thecostyle. The ovoid to elongate body protrudes to a greater or lesser extent beyond the apical rim 
of thecostyle although the latter is sufficiently large to enclose at least half of the zooid. Capitate 
tentacles not in fascicles, usually covering the apical surface of the exposed part of the zooid. Apical 
part of thecostyle is triangular, tapering posteriorly to form a hollow tube-like stem that is at least 
three times the length of the lorica-like part. The junction between the two parts of the thecostyle 
often, secondarily, narrowed and flexible. Attached to hydroid colonies and marine algae. 
Single contractile vacuole usually positioned laterally. Spherical macronucleus located centrally. 
Reproduction by exogenous budding resulting in an ovoid swarmer covered in transverse ciliary 
rows with some anterior short residual tentacles. 

NOTE. The observation by Collin (1912) that the formation of a narrow flexible junction between 
stem and lorica is a secondary event allows the inclusion of Acineta divisa Fraipont, 1 878 as a junior 
synonym. 

Paracineta irregularis Dons, 1928 

DESCRIPTION (Fig. 18). This is a small (15-25 um long), marine suctorian with a thecostyle. The 
ovoid to irregularly shaped body protrudes to a greater or lesser extent beyond the apical rim of 
thecostyle although the latter half of the zooid is always enclosed. Tentacles cover the apical 



C. R. CURDS 




Fig. 17 Paracineta patula: (a-c) after Collin, 1912; (d-e) after Claparede & Lachmann, 1861 (called 
Acineta patula); (f) after Fraipont, 1877 (called Acineta divisa); (g) after Calkins, 1902 (called Acineta 
divisd). 



surface of the exposed part of the zooid. Apical part of thecostyle irregularly triangular, tapering 
posteriorly to form a rigid hollow tube-like stem that is at least half the length of the lorica-like 
part. Epizoic on chaetae of the polychaete worm Pherusa plumosa. Spherical macronucleus located 
centrally. Reproduction not described. 

Paracineta jorisi (Sand, 1895) Collin, 1912 
AcinetajorisiSand, 1895 

DESCRIPTION (Fig. 19). This is a small to medium (30-80 urn long), marine suctorian with a 
thecostyle. The ovoid to pyriform body protrudes to a great extent beyond the apical rim of 
the semi-lorica part of the thecostyle which is not normally large enough to enclose the zooid. 
Tentacles not in fascicles, usually covering the apical surface of the exposed part of the body. 
Apical part of thecostyle is triangular or cup-like. The rim is prominently flared and folds back on 
itself to form an internal layer upon which the zooid is mounted. Thecostyle tapers posteriorly to 



PARACINETA AND CORYNOPHRYA PROBLEM 



89 




Fig. 18 Paracineta irregularis: (a-e) various forms after Dons, 1928. 




Fig. 19 Paracineta jorisi after Sand, 1 895 (called Acinetajorisi). 



form a rigid hollow tube-like stem that is at least three times the length of the lorica-like part. 
Attached to hydroid colonies such as Vesicularia and Sertularia. Single contractile vacuole. 
Spherical macronucleus located centrally. Reproduction by exogenous budding. 



90 



C. R. CURDS 




Fig. 20 



Paracineta limbata: (a) adult with swarmer, after Collin, 1912; (b) after Wailes, 1928; (c) after 
Dons, 1922; (d,e) after Moebius, 1888 (called Podophrya limbata). 



Paracineta limbata (Maupas, 1881) Collin, 1912 

Podophrya limbata Maupas, 1881 
Tokophrya limbata Biitschli, 1889 
Paracineta limbata forma convexa Dons, 1922 
Luxophrya limbata Jankowski, 1978 

DESCRIPTION (Fig. 20). This is a small (20-45 um diameter), marine suctorian with a thecostyle. 
The spherical body is mounted on the rim of a greatly reduced lorica-like part of the thecostyle. 
Zooid often covered by a thick gelatinous outer coat. Capitate tentacles not in fascicles, radiate out 
from the entire surface of the exposed zooid. Reduced apical part of thecostyle is cone-like, 
tapering posteriorly to join a rigid hollow tube-like stem that is at least four times the diameter of 
the zooid in length. Attached to hydroid colonies. Two contractile vacuoles usually positioned 
laterally. Spherical macronucleus located centrally. Reproduction by exogenous budding resulting 
in an ovoid swarmer covered in transverse ciliary rows with some residual tentacles. 



PARACINETA AND CORYNOPHRYA PROBLEM 



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Fig. 21 Paracineta moebiusi after Moebius, 1 888 (called Acineta crenatd). 



Paracineta moebiusi (Moebius, 1888)Kahl, 1934 
Acineta crenata Moebius, 1888 

DESCRIPTION (Fig. 21). This is a medium (76 um long), marine suctorian with a thecostyle. 
Approximately half the elongate body protrudes beyond the apical rim of thecostyle. Tentacles not 
in fascicles, covering only the apical surface of the exposed part of the zooid. The thecostyle is 
prominently and totally ribbed transversely. The apical part is cup-shaped, and tapers posteriorly 
to form a rigid hollow tube-like stem that is about one and a half times the length of the lorica- 
like part. Epizoic on the crustacean Holocarus. Single anterior contractile vacuole. Spherical 
macronucleus located posteriorly. Reproduction not described. 

Paracineta vorticelloides (Fraipont, 1877) Collin, 1912 

Acineta vorticelloides Fraipont, 1877 
Proluxophrya vorticelloides Jankowski, 1978 

DESCRIPTION (Fig. 22). This is a small (30-40 um diameter), marine suctorian with a thecostyle. 
The spherical body is mounted on the greatly reduced anterior part of the thecostyle. Capitate 
tentacles not in fascicles, radiating out from the entire surface of the exposed zooid. Reduced apical 
part of thecostyle is cup-like, tapering posteriorly to join a rigid hollow tube-like stem that is at 
least four times the diameter of the body in length. Epizoic on hydroid colonies, Crustacea and 
marine algae. Single central contractile vacuole. Spherical macronucleus located posteriorly. 
Reproduction by exogenous budding. 



Genus LORICOPHRYA Matthes, 1956 



Acineta Ehrenberg, \S33proparte 
Thecacineta Collin, \9Q9proparte 
Paracineta Collin, 191 1 pro par te 



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C. R. CURDS 




Fig. 22 Paracineta vorticelloides: (a,b) after Fraipont, 1878 (called Acineta vorticelloides). 



Corynacineta Jankowski, 1978 
Heliotheca Jankowski, 1978 
Paraloricophrya Jankowski, 1978 
Spongiophrya Jankowski, 1978 

The genus was originally erected by Matthes (1956) for loricate suctoria with a single apical 
group of tentacles but with an unknown method of budding. He designated Loricophrya parva 
(Schulz, 1932) as the type species and listed the following species to constitute the genus: 
Loricophrya cattanei (Parona, 1883), L. simplex (Maskell, 1886), L. lasanicola (Maskell, 1887), 
L. tulipa (Maskell, 1887), L. solenophryaformis (Sand, 1899), L. cypridinae (Collin, 1912), 
L. caepula (Penard, 1920), L. edmondsoni(Kmg, 1932), L. sivertseni (Allgen, 1951), L. trichophora 
(Allgen, 1951) and L. longe-petiolatus (Allgen, 1951). The present author does not consider all of 
these species to be congeneric although the majority are retained in this revision. The three species 
described by Maskell (1886, 1887) have already been transferred back (Curds, 1985) into the 
genus Acineta but the generic position of L. cattanei (Parona, 1883) is still uncertain. Similarly, 
L. cypridinae (Collin, 1912) will be returned back to its original genus Thecacineta. All the others 
in Matthes (1956) original list have been retained within the genus although the specific epithet 
may be different to that used by him and several additions have been made. 

Diagnosis of Loricophrya 

Freshwater or marine sectoria with a thecostyle. When clearly differentiated the stem is shorter 
than the lorica part of the thecostyle. Body ovoid to elongate, rounded in cross-section. Capitate 
tentacles restricted to a single group on the apical surface of the zooid. Mode of reproduction not 
yet recorded. 

Key to the species of Loricophrya 

1 Thecostyle continually narrows posteriorly without a stalk region being clearly differentiated . 2 
A narrow stalk region is clearly differentiated from the rest of the thecostyle .... 4 

2 Most of zooid projects out of short thecostyle L.ovifornus 

Most of zooid enclosed within long thecostyle 3 

3 Zooid small, pyriform, lying in apical quarter of thecostyle L.tuba 

Zooid large, elongate, filling most of thecostyle cavity L. sivertseni 



PARACINETA AND CORYNOPHRYA PROBLEM 



93 



10 




Fig. 23 Loricophrya parva: (a,b) after Schulz, 1932 (called Thecacinetaparvd). 



10 



Zooid longer than wide, never dorso-ventrally flattened nor discoidal. 
Zooid wider than length, flattened dorso-ventrally or discoidal in shape 

Lorica part of thecostyle striated transversely 

Lorica part of thecostyle without striations or ribs 

Thecostyle wider than height, covered in tubercles 

Thecostyle longer than wide, smooth 

Stalk part of thecostyle is half length of lorica part, and may be striated 

Stalk part of thecostyle very short, about 1 /8 of lorica part, not striated 

Stalk part of thecostyle striated, lorica part triangular in outline . 

Stalk part of thecostyle not striated, lorica part oval in outline . 

Stalk region very short, about 1/8 length of lorica region. Rim without collar, 

Stalk region short, about 1/2 length of lorica region. Rim of thecostyle 

surrounding wide aperture 

Stalk part of thecostyle conical in shape 

Stalk part of thecostyle tubular 



5 
9 

. L. lauterborni 
6 

. L. bifaria 
1 
8 

. L. multitentaculata 

. L. stresemanni 

. L. trichophora 

small aperture L. caepula 

with collar region 

. 10 

L. parva 

. L. solenophryaformis 



Species descriptions 

Loricophrya parva (Schulz, 1932) Matthes, 1956 
Thecacineta parva Schulz, 1932 

DESCRIPTION (Fig. 23). This the type species is a small (36-41 um long), brackish- water suctorian 
with a thecostyle. The discoidal body is rounded in cross-section and lies within an urn-like 
thecostyle. There is a single apical group of capitate tentacles on the apical surface. The 
thecostyle narrows somewhat posteriorly to form a cone-like stalk region. Attached to inanimate 
objects. Single lateral contractile vacuole. Macronucleus oval, centrally located. Reproduction 
not described. 

Loricophrya bifaria (Stokes, 1887) n. comb. 

Acineta bifaria Stokes, 1887 
Paracineta bifaria Collin, 1912 
Paraloricophrya bifaria Jankowski, 1978 



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Fig. 24 



Loricophrya bifaria: (a) adult; (b) budding; (c) swarmer; all after Stokes, 1887 (called Acineta 

bifarid). 




Fig. 25 Loricophrya caepula after Penard, 1920 (called Thecacineta caepuld). 



DESCRIPTION (Fig. 24). This is a small (45 urn diameter), freshwater suctorian with a thecostyle. The 
elongate body is rounded in cross-section and projects out well beyond the rim of the thecostyle. 
There is a single group of apical capitate tentacles. Stalk region a short, button-like projection. 
Lorica region ovoid, covered in tubercles, width greater than height. Single lateral contractile 
vacuble. Ovoid macronucleus centrally located. Reproduction by exogenous budding resulting in 
an elongate swarmer with longitudinal rows of cilia and some residual tentacles. 

Loricophrya caepula (Penard, 1920) Matthes, 1956 

Thecacineta caepula Penard, 1920 
Heliotheca caepula Jankowski, 1978 

DESCRIPTION (Fig. 25). This is a small (33 \im diameter), freshwater suctorian with a thecostyle. 
The ovoid body is rounded in cross-section and just projects out beyond the rim of the thecostyle. 
There is a single group of apical capitate tentacles. Stalk region a short, button-like projection. 
Lorica region ovoid, width greater than height. Single anterio-lateral contractile vacuole. Ovoid 
macronucleus centrally located. Reproduction not described. 



PARACINETA AND CORYNOPHRYA PROBLEM 



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Fig. '26 Loricophrya lauterborni: (a) after Sondheim, 1929 (called Paracineta lauterborni); (b) after 

King, 1932 (called Thecacineta edmondsi). 



Loricophrya lauterborni (Sondheim, 1929) n. comb. 

Paracineta lauterborni Sondheim, 1929 
Thecacineta edmondsi King, 1932 
Paraloricophrya lauterborni Jankowski, 1978 

DESCRIPTION (Fig. 26). This is a small (40-55 um diameter), freshwater suctorian with a thecostyle. 
The ovoid body is rounded in cross-section and projects out beyond the rim of the thecostyle. 
Capitate tentacles radiate out from the surface of the exposed part of the zooid. Stalk region a 
short, button-like projection or up to half the lorica length. Lorica region cup-like with about 
four transverse rings. Attached to inanimate objects. Two or three contractile vacuoles. Ovoid 
macronucleus centrally located. Reproduction possibly by exogenous budding. 

Loricophrya multitentaculata (Sand, 1895) n. comb. 

Hallezia multitentaculata Sand, 1895 
Acineta multitentaculata Sand, 1899 
Paracineta multitentaculata Collin, 1912 
Spongiophry a- multitentaculata Jankowski, 1978 

DESCRIPTION (Fig. 27). This is a large (304 um long), marine suctorian with a thecostyle. The 
cylindrical body is rounded in cross-section and only the small posterior part is housed in the 
cup-like thecostyle. There is a single apical group of capitate tentacles on the apical surface. The 
thecostyle follows the outline of the body and there is a short button-like stalk region. Epizoic on 
sponges such as Leucosolenia. Contractile vacuole not observed. Macronucleus large, elongate, 
centrally located. Reproduction not described. 



96 



C. R. CURDS 




Fig. 27 



Loricophrya multitentaculata after Sand, 1895 (called Hallezia multitentaculatd). Note that the 
theca was described but not illustrated in the original description. 



NOTE. The presence of a lorica was not shown in the diagram of this species but was mentioned 
clearly in the description. Here the presence of a lorica is indicated means of dotted lines. 

Loricophrya oviformis (Dons, 1918) n. comb. 
Par acineta oviformis Dons, 1918 

DESCRIPTION (Fig. 28). This is a medium (85 urn long), marine suctorian with a thecostyle. The 
ovoid body is only partially enclosed within the thecostyle whose rim is smooth. There is a single 
group of tentacles which are scattered over much of the exposed body surface. The thecostyle 
follows the outline of the body posterior and there is a short button-like stalk-region. Epizoic on 
the worm Spirorbis. Nuclear and reproductive features not described. 

Loricophrya sivertseni (Allgen, 1951) Matthes, 1956 
Thecacineta sivertseni Allgen, 1951 

DESCRIPTION (Fig. 29). This is a large (108um long), marine suctorian with a thecostyle. The 
elongate body is totally enclosed within the cone-shaped thecostyle whose rim is scalloped. 




Fig. 28 Loricophrya oviformis after Dons, 1918 (called Paracineta oviformis). 



PARACINETA AND CORYNOPHRYA PROBLEM 



97 




10 



Fig. 29 Loricophrya sivertseni after Allgen, 1951 (called Thecacineta sivertseni). 




Fig. 30 Loricophrya solenophryaformis after Sand, 1 899 (called Acineta solenophryaformis). 



Capitate tentacles in a single apical group. There is no distinct stalk region, the lorica gradually 
and continually narrows posteriorly to join the attachment plate. Epizoic on the nematode worm 
Spirina parasitifera. Ovoid macronucleus centrally located. Mode of reproduction not described. 

Loricophrya solenophryaformis (Sand, 1899) Matthes, 1956 

Acineta solenophryaformis Sand, 1899 
Thecacineta solenophryaformis Collin, 1909 

DESCRIPTION (Fig. 30). This is a small (30-35 um long), freshwater suctorian with a thecostyle. The 
discoid body is totally enclosed within an urn-like thecostyle whose rim is surrounded by a collar- 
like region. Capitate tentacles located in a single, tightly-packed, apical group which are enclosed 
within the thecostyle. There is a short but distinct, tubular stalk region. Attached to freshwater 
algae. Ovoid macronucleus located posteriorly. Mode of reproduction not described. 

Loricophrya stresemanni (Allgen, 1951) Matthes, 1956 
Paracineta stresemanni Allgen, 1951 
DESCRIPTION (Fig. 31). This is a small (40 urn long), marine suctorian with a thecostyle. The 



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C. R. CURDS 





Fig. 31 Loricophrya stresemanni: (a,b) after Allgen, 1951 (called Paracineta stresemanni). 



elongate body is mostly enclosed within a cone-shaped thecostyle whose rim is smooth. Capitate 
tentacles in a single apical group. There is a distinct stalk region which is about half the lorica 
length and is striated transversely. Epizoic on the nematode worm Spirina parasitifera. Ovoid 
macronucleus centrally located. Mode of reproduction not described. 

Loricophrya trichophora (Allgen, 1951) Matthes, 1956 

Thecacineta trichophora Allgen, 1951 
Thecacineta longe-petiolatus Allgen, 1951 

DESCRIPTION (Fig. 32). This is a medium (80 um long), marine suctorian with a thecostyle. 
The elongate body is totally enclosed within an ovoid thecostyle whose rim is smooth. Capitate 
tentacles in a single apical group. There is a distinct stalk region which is about half the lorica 
length, not striated. Epizoic on the nematode worm Spirina parasitifera. Ovoid macronucleus 
centrally located. Mode of reproduction not described. 

Loricophrya tuba (Zelinka, 1914) n. comb. 

Acineta tuba Zelinka, 1914 
Paracineta tuba Kahl, 1934 
Corynacineta tuba Jankowski, 1978 

DESCRIPTION (Fig. 33). This is a small (25-32 um long), marine suctorian with a thecostyle. The 
pyriform body is enclosed within the apical quarter of the elongated cone-like thecostyle. Tentacles 
emerge from the apical surface, not in fascicles. There is no distinct stalk region, the lorica 
gradually and continually narrows posteriorly to join the substratum. Epizoic on the shells of 
echinoderms. Ovoid macronucleus centrally located. Mode of reproduction not described. 

Genus ANTH ACINETA Jankowski, 1978 

Acineta Ehrenberg, 1833 pro pane 
Noracine ta Jankowski, 1978 



PARACINETA AND CORYNOPHRYA PROBLEM 



99 




10 



Fig. 32 Loricophrya trichophora after Allgen, 1951 (called Paracineta trichophord). 



10 




Fig. 33 Loricophrya tuba after Zelinka, 1914 (called Acineta tuba). 



The genus Anthacineta was erected by Jankowski (1978) for Acineta craterellus Collin, 1909 giving 
the following brief diagnosis 'semi-lorica - stylotheca'. According to that brief definition the genus 
could be transferred to Paracineta and several other similar genera as a junior synonym. It can only 
be classified as a distinct genus if the two fascicles of tentacles and rounded transverse section to the 
body are taken into account. Here the diagnosis has been expanded and one other species, Acineta 
infundibuliformis Wang & Nie, 1933, has been transferred to it for the first time. 

Diagnosis of Anthacineta 

Marine suctorians with thecostyle. Zooid only partly enclosed in the semi-lorica part of the 
thecostyle which has a long stem. Body ovoid, rounded in cross-section. Two fascicles of capitate 
tentacles present, one either side of the zooid. Mode of reproduction not recorded. 



100 



C. R. CURDS 




JO. 



Fig. 34 Anthacineta craterellus after Collin, 1912 (called Acineta craterellus). 



Key to the species of Anthacineta 

1 Zooid wider than long, contractile vacuole in posterior body half. Thecostyle narrows abruptly to form 

stalk-like region . A. infundibuliformis 

Zooid longer than wide, contractile vacuole in anterior body half. Thecostyle narrows consistently to 
form the stalk-like region 5. craterellus 



Species descriptions 

Anthacineta craterellus (Collin, 1909) Jankowski, 1978 

Acineta tuber osa Sand, 1901 propane 
Acineta craterellus Collin, 1909 

DESCRIPTION (Fig. 34). This the type species is a small (50 ^m long), marine suctorian with a 
thecostyle. The ovoid to pyriform body is rounded in cross-section and about half of it protrudes 
beyond the rim of the semi-lorica part of the thecostyle. There are two anterio-lateral fascicles 
of capitate tentacles. The lorica part of the thecostyle is short and cone-like, narrowing gently 
posteriorly to form the hollow stem region that is at least twice the length of the zooid. Epizoic on 
bryozoa. Single contractile vacuole situated apically between fascicles. Spherical macronucleus 
centrally located. Reproduction not described. 

Anthacineta infundibuliformis (Wang & Nie, 1933) n. comb. 

Acineta infundibuliformis Wang & Nie, 1933 
Noracineta infundibuliformis Jankowski, 1978 

DESCRIPTION (Fig. 35). This is a small (50 um long), marine suctorian with a thecostyle. The 
wedge-shaped body is rounded in cross-section and about half of it protrudes beyond the rim of the 
semi-lorica part of the thecostyle. There are two lateral fascicles of capitate tentacles. The lorica 
part of the thecostyle is short and cone-like, narrowing abruptly posteriorly to form the hollow 
stem region that is about the length of the lorica. Attached to marine algae. Single contractile 
vacuole situated posteriorly. Ovoid macronucleus centrally located. Reproduction not described. 



PARACINETA AND CORYNOPHRYA PROBLEM 



101 




10 



Fig. 35 Anthacineta infundibuliformis after Wang & Nie, 1933 (called Acineta infundibuliformis). 



Genus FLECTACINETA Jankowski, 1978 

Acineta Ehrenberg, \S33proparte 
Podophrya Ehrenberg, 1833 pro par te 
Alderia Alder, 1851 
Paracineta Collin, 191 1 pro par te 

The genus was erected by Jankowski (1978) for Acineta livadiana Mereschowsky, 1881 who gave 
the following brief diagnosis, 'with stylotheca and apical tentacles'. The stalk is normally shown as 
being hollow but not as an extension of the lorica as the term stylotheca implies. Thus the diagnosis 
has been emended slightly and expanded for the sake of clarity. Two species Paracineta dadyi 
(Daday, 1886) Kahl, 1934 and Acineta elegans Imhoff, 1883 have been transferred to the genus for 
the first time. 

Diagnosis of Flectacineta 

Marine loricate suctorians. Ovoid body, rounded in cross-section lying within lorica. Capitate 
tentacles restricted to single apical group. Thecostyle lorica rim characteristically inverted at apex, 
mounted upon a hollow stalk. Reproduction by exogenous budding. 



Key to the species of Flectacineta 

1 Rim of lorica smooth, junction between stalk and lorica simple . 
Rim of lorica scalloped, junction between stalk and lorica complex 

2 Wall or lorica divided into an inner and outer wall near aperture . 
Wall of lorica not divided . 



. F. elegans 

F. dadyi 

F. livadiana 



Species descriptions 
Flectacineta livadiana (Mereschkowsky, 1881) Jankowski, 1978 



Cothurnia havniensis Ehrenberg, 1838 
Alderia pyriformis Alder, 1851 
Podophrya pyriformis Pritchard, 1861 
Acineta livadiana Mereschkowsky, 1881 
Acineta neapolitana Daday, 1886 
Acineta sp. Robin, 1879 
Paracineta neapolitana Kahl, 1934 



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Fig. 36 Flectacineta livadiana: (a) after Sand, 1895 (called Acineta livadiana); after Mereschkowsky, 
1881 (called Acineta livadiana); (c) after Wang & Nie, 1933 (called Acineta livadiana); (d) after Daday, 
1886 (called Acineta neapolitana). 



DESCRIPTION (Fig. 36). This the type species is a small to medium (30-80 um long), marine, loricate 
suctorian. The small ovoid body is rounded in cross-section and is completely enclosed within the 
lorica. There is a single apical group of capitate tentacles. The lorica is ovoid with an inverted rim 
that forms a small aperture. The stem region is distinct and most diagrams show that there is 
usually at least a narrow channel through the centre. Length of stem variable. Epizoic on hydroids 
and marine algae. Single contractile vacuole situated laterally. Ovoid macronucleus centrally 
located. Reproduction by exogenous budding. 

Flectacineta dadayi (Daday, 1886) n. comb. 

Acineta livadiana Daday, 1886 
Paracineta livadiana Collin, 1912 pro pane 
ParacinetadadayiKahl, 1934 

DESCRIPTION (Fig. 37). This is a small (45 urn long), marine, loricate suctorian. The small ovoid 
body is rounded in cross-section and is completely enclosed within the lorica. There is a single 
apical group of capitate tentacles. The lorica is ovoid to cone-shaped with an inverted rim that 
forms a small aperture. The lorica surrounding the aperture is divided into an inner and an 
outer wall. The hollow stem region is distinct and some diagrams show that there is a narrow 



PARACINETA AND CORYNOPHRYA PROBLEM 



103 




Fig. 37 Flectadneta dadayi: (a) after Daday, 1886 (called Acineta livadiana); (b) after Collin, 1912 

(called Paracineta livadiana}. 




Fig. 38 Flectadneta elegans after Imhoff, 1 884 (called Acineta elegans). 

channel through the centre. Length of stem region about that of lorica. Epizoic on hydroids and 
marine algae. Single contractile vacuole situated laterally. Ovoid macronucleus centrally located. 
Reproduction not described. 



104 C. R. CURDS 

Flectacineta elegans (Imhoff, 1883) n. comb. 

Acineta elegans Imhoff, 1883 non Maskell, 1886 
Paracineta elegans Collin, 1912 

DESCRIPTION (Fig. 38). This is a medium (70 jim long), marine, loricate suctorian. The rectangular 
body is rounded in cross-section and is completely enclosed within the lorica. There is a single 
apical group of capitate tentacles. The lorica is pyriform with an inverted scalloped rim that 
forms a small aperture. The hollow stem region is distinct and joins the lorica via an intervening 
ball-like joint. Length of stem at least twice that of the lorica. Epizoic on the cladoceran 
Bythotrephes longimanus. Single apical contractile vacuole. Ovoid macronucleus centrally located. 
Reproduction not described. 



References 

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Fraipont, T. 1877. Recherches sur les Acinetiens de la cote d'Ostende. Bulletin de I' Academic Royale de 

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Imhoff, O. E. 1883. Studien zur Kentniss der pelagischen Faunen der Schweizeseen. Zoologischer Anzeiger. 

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1 884. Resultate meiner Studien uber die pelagische Fauna kleinerer und grosserer Susswasserbecken der 

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Fortpflanzung durch Vermoid-Schwarmer. Archivfur Prostistenkunde 101: 477-528. 
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9: 299-368. 
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153-248. 

1881. On some new or little-known Infusoria. Annals and Magazine of Natural History; including 



Zoology, Botany and Geology. London 7 (5th Ser): 209-219. 
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1888:81-116. 
Parana, C. 1881. Delle Acinetine in generale ed in particulare di una nouva forma (Acineta dibdalteria n.sp.) 

Bollettino Scientifico. Milano 1880: 79-85. 
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Akademieder Wissenschaften. Mathematisch-Naturwissenschaftliche Classe. Wien 120: 757-763. 
Schulz, E. 1932. Beitrage zur Kenntniss mariner Suctorien. Zoologischer Anzeiger. Leipzig 97: 289-292. 

1933. Beitrage zur Kenntnis mariner Suctorien. Zoologischer Anzeiger. Leipzig 103: 327-329. 

Sewell, R. B. S. 1951. The epibionts and parasites of the Planktonic Copepoda of the Arabian Sea. Scientific 

Reports. The John Murray Expedition 1933-34. London 39: 255-394. 
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Abhandlungen der Senckenbergische Naturforschende Gessellschafte 41: 283-313. 



106 



C. R. CURDS 



Steuer, A. 1928. Uber eine neue Paracineta aus dem Sudatlantik. Sitzungsberichte der Akademie der Wissen- 

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Gesellschaft Deutscher Naturforscher. Leipzig 85: 680-683. 

Manuscript accepted for publication 7 April 1986 



Index to Species 

Names given in roman refer to synonyms 



Acineta bifaria 93 

conipes 79 

craterellus 100 

crenata Fraipont 73 

crenata Moebius 90 

dibdalteria 84 

divisa 87 

elegans 104 

euchaetae 84 

homari 75 

infundibuliformis 100 

jorisi 88 

livadiana Daday 102 

livadiana Mereschkowsky 101 

lyngbyi 78 

multitentaculata 95 

neopolitana 101 

patula 87 

sai fulae 73 

solenophryaformis 97 

sp. Robin 101 

tuba 98 

tuberosa 100 

vorticelloides91 
Actinocyathula cidaris 73 

crenata 73 

gaetani 75 

homari 75 

pleuromammae 76 
Actinocyathus cidaris 73 
Alderia pyriformis 101 
Anthacineta craterellus 100 

infundibuliformis 100 

Corynophrya campanula 83 
columbiae 79 
conipes 79 
crenata 73 
francottei 79 
homari 75 
interrupta 83 
lyngbyi 78 
macropus 8 1 
steueri 83 
symbiotica%\ 
tuba 98 



tumida 8 1 
Cothurnia havniensis 101 

Discophrya campanula 83 
francottei 79 
interrupta 83 
lyngbyei 78 
steueri 83 
tumida 8 1 

Ephelota columbiae 79 

Faltacineta gataeni 75 

pleuromammae 76 
Flectacineta dadayi 102 

elegans 104 

livadiana 101 

Hallezia multitentaculata 95 
Heliotheca caepula 94 

Loricophrya bifaria 93 

caepula 94 

lauterborni 95 

multitentaculata 95 

oviformis 96 

parva 93 

sivertseni 96 

solenophryaformis 97 

stresemanni 97 

trichophora 98 

tuba 98 
Luxophrya limbata 90 

Miracineta saifulae 73 
Noracineta infundibuliformis 100 

Paracineta bifaria 93 
crenata 73 

crenata forma pachytheca 73 
crenata var. pachytheca 73 
dadayi 102 
divisa 87 
elegans 104 
gataeni 75 



homari 75 

irregularis 87 

jorisi 88 

lauterborni 95 

limbata 90 

limbata forma convexa 90 

livadiana 102 

moebiusi9\ 

multitentaculata 95 

neapolitana 101 

oviformis 96 

patula 87 

pleuromammae 76 

stresemanni 97 

tuba 98 

vorticelloides 91 
Paraloricophrya bifaria 93 

lauterborni 95 
Pelagacineta campanula 83 

dibdalteria 84 

euchaetae 84 

interrupta 83 
Podophrya conipes 79 

limbata 88 

lyngbyei 78 

macropus 81 

pyriformis 101 
Proluxophrya vorticelloides 91 

Spongiophrya multitentaculata 95 
Stemacineta patula 87 

Thecacineta caepula 94 

edmondsi 95 

longe-petiolatus 98 

parva 93 

sivertseni 96 

solenophryaformis 97 

trichophora 98 
Tokophrya campanula 83 

conipes 79 

francottei 79 

interrupta 83 

limbata 88 

lyngbyei 78 

steueri 83 




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1980, Hardcovers, 222pp, 17.50 (18.00 by post) Co-published by British Museum (Natural History) and 
Oxford University Press. 

This book illustrates, using scanning electron micrographs, most of the common species of testate amoebae 
that are found in freshwater habitats. Information on the biology, ecology, geographical distribution and a 
classification are followed by descriptions of ninety-five species. Each of these is illustrated by several views 
of the shell. The text if designed not only to enable biologists to identify species of testate amoebae, but to 
serve as an introduction to students interested in the taxonomy and biology of these freshwater protozoa. It 
will be of special interest to protozoologists, ecologists, limnologists, water treatment specialists and 
micropalaeontologists interested in recent sediments. 

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Notes on spiders of the family Salticidae 1. The genera Spartaeus, Mintonia and 
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Mites of the genus Holoparasitm Oudemans, 1936 (Mesostigmata: Parasitidae) in 
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The phylogenetic position of the Yugoslavian cyprinid fish genus Aulopyge Heckel, 
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Notes on spiders of the family Salticidae. 
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F. R. Wanless 



Zoology series Vol52 No 3 26 March 1987 



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Notes on spiders of the family Salticida^. 

Ir^.1 o J. . *jj ^ 2 6nAS.i7o7 

. 1 he genera bpart aeus, Mintonia and; Taraxi 



F. R. Wanless 

Department of Zoology, British Museum (Natural History), 

"X^ ^ffAl Hl^> ' 

&^ 

I 

Synopsis 

The genus Taraxella is redefined to include those spartaeines in which the embolus of the male palp is largely 
obscured by tegular apophyses. The male of Spartaeus thailandica Wanless and the female of Mintonia 
melinauensis Wanless are described for the first time. One new species of Spartaeus, two new species of 
Mintonia and four new species of Taraxella are described from the Oriental Region. Diagnoses and figures are 
provided. The presence of mytiliform organs on the legs and filamentous metatarsal leg fringes are described 
for the first time. They are illustrated, together with femoral organs, muscle attachment sites and apophyses 
by scanning electron micrographs. 



Introduction 

The purpose of the present paper is to describe seven new species and the previously unknown male 
and female of two described species belonging in the subfamily Spartaeinae. The subfamily is of 
particular interest because some species spin large webs that are used to capture prey. Typical 
salticids are cursorial hunters with good vision that do not spin webs to capture prey, although they 
will spin silk nests in which to lay eggs, moult and sometimes mate, and generally rest at night or 
during other periods of inactivity (Jackson, 1979). 

The first reports of web-spinning in jumping spiders (Coleman, 1978, 1980; Murphy, in Wanless, 
19786) were followed by several important studies (Jackson & Blest, 1982; Jackson, 1982; Jackson 
& Hallas, in press a) that confirmed the phenomenon and provided a rare insight into the biology of 
a small group of tropical salticids. These spiders all belonged in the old world genus, Portia Karsch, 
that is presently classified along with 12 other genera in the subfamily Spartaeinae. In addition to 
building large prey-capture webs, Portia species will leave their web and stalk prey as cursorial 
hunters i.e. in the same manner as other salticids. Furthermore, they may invade the webs of other 
spiders and feed on trapped insects (kleptoparasitism), the resident spider, or even its eggs 
(oophagy). In life they resemble tatty mouldy leaves or detritus, their ornate hair tufts and fringes 
(Fig. 3) providing a form of camouflage that enables them to stalk prey without being noticed, an 
important guise since they show a marked preference for other spiders, including salticids. Portia 
species are also 'aggressive vibratory mimics' for when they invade other spider webs they pluck 
the threads and deceive the owner into accepting Portia as potential prey only to be attacked 
themselves on approaching within jumping distance. 

Occurrence of the unusual behaviour patterns of Portia species correlate to some degree in this 
and related spartaeines by the presence of morphological structures that are not known to occur in 
other spiders. These include femoral organs (Figs 5E; 14A) and pore-bearing apophyses (Figs 9E 
arrowed; 20 A, B) both of which are especially evident in some of the species described below. Also 
present on the legs of one species (Spartaeus wildtrackii sp. n.) are mytiliform organs (Fig. 15A-C; 
16A, B), structures that have hitherto only been found grouped together as a discrete patch on the 
dorsal surface of the abdomen (Fig. 20C) of species ofCyrba Simon, Portia, Gelotia Thorell, and 
Mintonia Wanless. The function of these structures is unknown but previous studies (Wanless, 
1984a, b; 1985) have suggested that they may be associated with pheromone dispersal. Jackson & 
Hallas (in press b} have demonstrated that sex pheromones are involved in mate recognition in 
some Portia, Brettus Thorell and Cyrba, thus supporting earlier work by Legendre and Llinares, 



Bull. Br. Mus. not. Hist. (Zool.) 52(3): 107-137 Issued 26 March 1987 

107 



108 



F. R. WANLESS 




Fig. 1. (above) Spartaeus wildtrackii sp. n. Subadult <$, under web on surface of tree trunk. Fig. 2 

(below) web of Spartaeus wildtrackii sp. n. 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 



109 




Fig. 3. Portia labiata (Thorell), 9 from Malaysia. 

1970 who noted that in Cyrba algerina (Lucas) pheromones left by the female stimulate the male 
and elicit searching behaviour. 

Another feature clearly demonstrated by some of the species described below is the disparity in 
size of the posterior median eyes. In most spartaeines and also in a few other subfamilies the 
posterior median eyes are relatively large (Fig. 5A, B) and fully functional, whereas in most 
jumping spiders they are relatively small (Fig. 10A, B) with no demonstrable function. It is 
therefore of interest to note that in the species of Taraxella Wanless, described below there is a clear 
trend towards the development of small posterior median eyes. In practice the distinction between 
relatively large or relatively small becomes somewhat blurred in Taraxella, although there has 
hitherto been no difficulty in assigning one state or the other. 

The presence of large posterior median eyes and web spinning behaviour in Portia species gave 
rise to the hypothesis (Jackson & Blest, 1982) that the ancestors of modern day salticids evolved 
from web building spiders with poorly developed vision and that acute vision, evolved originally in 
a spider like Portia that became an araneophagic predator, proficient at invading diverse types of 
webs. Subsequent papers on morphology (Wanless, 19840, b) the evolution of salticid eyes (Blest, 
1984; Blest & Sigmund, 1985) and behaviour (Jackson, 19856; Jackson & Hallas, in press a, b) 
have given support to the hypothesis and suggest that although spartaeine salticids are highly 
specialized they may nevertheless represent one of the most primitive branches of the family. 

Studies, however, are at an early stage for the biology of tropical salticids is very poorly known, 
in fact the majority of species cannot even be identified with confidence. Also, it is important to 
appreciate that the subfamily Spartaeinae, with more than 60 described species, represents less 
than 1-5% of the worlds salticid fauna. Despite this there are indications that jumping spiders 
make far more use of silk than has hitherto been supposed, for recent studies on other salticid 
groups have revealed species that build large prey-capture webs (Jackson, 19850) and even species 
that live in groups forming nest complexes within the webs of other spiders (Jackson, in press). 



110 F. R. WANLESS 

The standard abbreviations and measurements are those made by Wanless (1978) but for the leg 
spination the system adopted is that used by Platnick & Shadab ( 1 975). Note also, that the covering 
hairs on the male palps are not shown in any of the figures, because they are usually rather dense 
and obscure details. 

Genus SPA R TA BUS Thorell 

Spartaeus: Wanless 1984a: 147 [synonymy, definition and species descriptions]. Blest & Sigmund, 1985: 129. 
Blest, 1985:96. 

Spartaeus is a small oriental genus comprising three species, S. spinimanus Thorell, from Indonesia, 
Malaysia and Sri Lanka, S. thailandicus Wanless from Thailand and S. wildtrackii sp. n., from 
Malaysia. They are easily distinguished by the structure of the genitalia. 

All three species possess relatively large posterior median eyes (Fig. 4A), unusually long slender 
legs bearing numerous spines, and well developed femoral organs (Fig. 4E; 5E; 14A-C) on the first 
pair of legs of adult males. Also present on the legs of both males and females are disc-like 
mytiliform organs (Figs 15C; 16B). Those on the femora are more or less rounded and sparsely 
distributed (Fig. 16A), whereas those on the tibiae are ovoid and located distally on the dorsal 
surface of the segment (Fig. 15A, B). As mentioned above, mytiliform organs have hitherto only 
been found in the form of a patch on the dorsal surface of the abdomen. Their occurrence on legs is 
therefore of particular interest, especially as some are grouped on the distal end of the tibiae, an 
arrangement that may be unique to Spartaeus. However, at present these structures cannot be 
used to determine relationships as their distribution on the legs of other salticids is unknown. 
Furthermore, they are almost certainly homologous with pustuliform organs (see Hill, 1977 and 
Wanless, 1984) that occur as scattered pore-bearing pustules on the abdomen, legs and pedipalps 
of Icius Simon, Metaphidippus F.O.P-C and Phidippus Koch, and as a group on the abdomen of 
Holcolaetis Simon and Sonoita Peckham and Peckham. 

The dorsal surface of the tibiae are characterised by hinge lines or rows of muscle attachment 
sites (Fig. 15A, B) that are electron dense and smooth in contrast to the surrounding cuticle. They 
resemble mytiliform organs, but lack pores and raised rims. It is also worth noting that they differ 
from the rows of triangular muscle attachment sites found on the tibiae and other leg segments of 
Holcolaetis species (see Wanless, 1985). 

The biology of Spartaeus is unknown except for observations (see below) made by Mr P. D. 
Hillyard (BMNH) who also provided the photographs (Fig. 1 ; 2) showing S. wildtrackii beneath its 
sheet web on a tree trunk. 

Spartaeus thailandicus Wanless 
(Fig. 4A-I) 

Spartaeus thailandica Wanless, 1984a: 151. Holotype ?, Thailand, BMNH, [examined]. 

DIAGNOSIS. 5". thailandicus seems to be most closely related to S. wildtrackii sp. n., but may be 
distinguished by the presence of a tegular apophysis (Fig. 4G, arrow) and the form of the retro- 
lateral tibial apophysis (Fig. 4F) in males; females are separated by the absence of a postepigynal 
furrow (see Wanless \9S4a, Fig. 5D). 

MALE, in good condition, from Khas Yai National Park, Thailand. Carapace (Fig. 4A-D): weakly 
iridescent under some angles of illumination; orange-brown lightly tinged and mottled black with a 
broad tapering pale yellow band on thoracic part; rubbed except for some black and pale amber 
hairs on sides. Eyes: laterals with black surrounds; fringed by white hairs. Clypeus: yellow-brown 
with sooty markings; sparsely clothed in black and whitish hairs. Chelicerae: long robust and 
diverging; posterior surface with series of transverse furrows; orange-brown lightly tinged black; 
sparsely clothed in black hairs with dense promarginal scopula; fang robust and curved; fang 
groove with eight promarginal teeth and 12 retromarginal denticles. Maxillae and labium: pale 
brownish yellow faintly tinged with some black. Sternum (Fig. 4B): pale yellow-brown with darker 
margins. Coxae: pale yellow-brown except for blackish sides of I and II. Abdomen: pale yellow- 
brown with black markings, ventrally a broad sooty band from epigastric furrow to spinnerets; 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 



111 




Fig. 4. Spartaeus thailandicus Wanless , A, dorsal view; B, sternum; C, chelicerae, maxillae and 
labium; D, carapace, lateral view; E, leg I; F, palp, retrolateral view; G, palp, ventral view; H, cheliceral 
teeth; I, palpal tibia, dorsal view. Abbreviations: e, embolus; fo, femoral organ; Ml, fan-shaped 
element of distal haematodocha. 



112 F. R. WANLESS 

rubbed; spinnerets long and robust. Legs: very long and slender, femoral organ pronounced; legs I 
yellow-brown except for blackish streaks on sides of femora, a blackish tinge towards apices of 
metatarsi and whitish yellow tarsi; other legs whitish yellow to yellow-brown with apices of 
metatarsi and tarsi tinged black; clothed in scattered simple and feathery hairs mostly rubbed; 
tarsi I-II and apices of metatarsi I with pro ventral row of specialised prey-capture setae; metatarsi 
II-III with basal fringe of fine curved filamentous setae. Spines strong and numerous; spination of 
leg I: metatarsus v 2^M), tibia v 4-6-7, femur p 1-1-1, d 0-2-2, r 0-1-1. Palp (Fig. 4F, G, I): 
element M 1 of the distal haematodocha (see Wanless, 19840) fan-shaped (Fig. 4G); element M2 is 
obscure and appears to have fused with the base of the embolus. 

Dimensions (mm): total length 8-5; carapace length 3-84, breadth 3-14, height 2-75; abdomen 
length 4-6; eyes, anterior row 2-7, middle row 2-04, posterior row 2-32; quadrangle length 1-9 
(49% of carapace length). 



Leg 


1 


2 


3 


4 


Palp 


Femur 


6-40 


3-84 


3-56 


4-56 


1-60 


Patella 


2-60 


1-72 


1-44 


1-60 


0-88 


Tibia 


7-04 


3-32 


3-16 


4-28 


0-56 


Metatarsus 


5-36 


3-16 


3-64 


5-28 






Tarsus 


1-96 


1-14 


1-16 


1-36 


1-68 



Total 23-36 13-18 12-96 17-08 4-72 



Ratios: AM : AL : PM : PL :: 22 : 13 : 8-5 : 12; AL PM PL : 12-14; AM : CL :: 22 : 5. 
DISTRIBUTION. Thailand. 

MATERIAL EXAMINED. Thailand: Khas Yai National Park, tropical evergreen forest, 1^ under bark of 
decomposing log, 17.iii.1984, P. D. Hillyard, BMNH. 1985.8.16.1. 

NATURAL HISTORY. The male described above was found together with several harvestmen 
(Opiliones) under the bark of a fallen decomposing log. There was no evidence of a web, but it was 
noticeable that the legs of this species were comparatively much longer than those of 5". wildtrackii, 
a character that may enable future collectors to distinguish the species in the field. 



Spartaeus wildtrackii sp. n. 
(Figs 1; 2; 5A-E; 6A-D; 14A-C; 15A&C; 16A-D; 17A-G; 18A-D) 

DIAGNOSIS. S. wildtrackii seems to be most closely related to S. thailandica, but may be dis- 
tinguished by the absence of a tegular apophysis and the form of the retrolateral tibial apophysis in 
males (Fig. 6A, B, D), and the presence of a postepigynal furrow in females (Fig. 5D). 

FEMALE HOLOTYPE, in fair condition. Carapace (Fig. 5A, B): weakly iridescent under some angles of 
illumination; orange-brown lightly tinged and mottled black with a broad yellow-brown tapering 
band on thoracic part and vague patches on sides; irregularly clothed in whitish hairs (mostly 
rubbed). Eyes: laterals with black surrounds; fringed by whitish and pale yellow hairs. Clypeus: 
sparsely clothed in black hairs. Chelicerae: robust, moderately long and more or less parallel; 
brownish orange lightly tinged black; thinly clothed in black hairs with dense promarginal scopula; 
promargin with six teeth, retromargin with nine or 12 denticles. Maxillae andlabium: pale orange- 
brown lightly tinged with some grey. Sternum: pale yellow with darker margins. Coxae: pale yellow 
except first pair with blackish promarginal sides. Abdomen: pale yellow with blackish markings and 
scattered clumps of whitish guanin; ventrally a broad black band from epigyne to spinnerets; 
spinnerets long and robust. Legs: long and slender; legs I pale yellow to light orange-brown with 
vague sooty markings and a proventral black stripe on femora that appears iridescent green under 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 



pm 




Fig. 5. Spartaeus wildtrackii sp. n., holotype <$: A, dorsal view; B, carapace, lateral view; D, epigyne. 
Paratype <$: C, sternum; E, leg, I. Paratype $: F, vulva, ventral view; G, vulva, dorsal view. 
Abbreviations: pe, postepigynal furrow; pm, posterior median eye; fo, femoral organ. 



some angles of illumination; other legs similar except femoral stripe lacking; also, dorsal spines 
arise from black spots; sparsely clothed in simple and feathery hairs, (Fig. 16D), mostly rubbed, 
with pro ventral row of specialized prey-capture setae on tarsi I-II (Fig. 17B, G) and apices of 
metatarsi I; proximal half of metatarsi II-III with scanty ventral fringe of fine curved filamentous 
setae (Fig. 1 7 A, C-F). Spines strong and numerous; spination of leg I: metatarsus v 3-2-0, p 0-1-0, 
tibia v 5-8-7, femur d 0-2-1. Palp: with terminal claw; pale yellow with greyish patch on tarsus; 
clothed in pale yellow and light greyish hairs. 



114 F. R. WANLESS 

Dimensions (mm): total length 6-4; carapace length 2-72, breadth 2-28, height 1-52; abdomen 
length 3-68; eyes, anterior row 2-08, middle row 1-48, posterior row 1-75; quadrangle length 1-58 
(58% of carapace length). 



Leg 


1 


2 


3 


4 


Palp 


Femur 


3-24 


2-56 


2-56 


3-16 


1-06 


Patella 


1-50 


1-16 


1-00 


1-06 


0-60 


Tibia 


3-16 


2-20 


2-20 


3-04 


0-64 


Metatarsus 


1-92 


1-74 


2-36 


3-20 






Tarsus 


0-84 


0-78 


0-92 


1-08 


1-16 


Total 


10-66 


8-44 


9-04 


11-54 


3-46 


: PM : PL :: 


16-5:11:7: 


11; AL- 


-PM- 


-PL :: 10-10; AM 



MALE PARATYPE, in good condition. Carapace: dark brown with dull orange-brown eye region and 
yellow-brown markings on thoracic part; clothed in fine recumbent light greyish and pale amber 
hairs. Eyes: laterals with black surrounds; fringed by pale yellow, whitish and amber hairs; also, a 
dense matt of short hairs behind anterior medians. Clypeus: tinged black, sparsely clothed in black 
hairs. Chelicerae: long, robust and slightly diverging; dark brown heavily tinged black; shiny; 
thinly clothed in black hairs with dense promarginal scopula; fang robust and curved with basal 
protuberance; promargin with eight teeth, retromargin with 12 (Fig. 6C). Maxillae and labium: 
orange-brown to yellow-brown tinged grey. Sternum (Fig. 5C): pale greenish yellow with darker 
margins; thinly clothed in greyish simple and feathery hairs. Coxae: pale greenish yellow with black 
promarginal stripe. Abdomen: yellow-brown suffused and mottled black; clothed in light and dark 
amber hairs with two spots comprised of whitish guanin; venter yellow-brown with grey-black 
band clothed in black feathery hairs from epigastric furrow to spinnerets; spinnerets long and 
robust. Legs: long and slender; femoral organ well developed (Fig. 5E; 14A-C); legs I pale greenish 
yellow to light orange brown with sooty markings and blackish longitudinal stripes on femora that 
shine iridescent green under some angles of illumination; thinly clothed in simple and black 
feathery hairs with some whitish ones on tarsi; specialized prey capture and filamentous hairs as 
in female. Spination of leg I: metatarsus v 3-1-1, p 0-0-1, r 1-0-0; tibia v 5-7-6; femur d 0-2-1, 
p 0-1-1. Palp (Fig. 6A, B, D): yellow-brown to light orange-brown with black iridescent stripe on 
underside of femora. 

Dimensions (mm): total length 6-96; carapace length 3-36, breadth 2-68, height 1-92; abdomen 
length 3-8; eyes, anterior row 2-32, middle row 1-62, posterior row 1-96; quadrangle length 1-8 
(54% of carapace length). 



Leg 


1 


2 


3 


4 


Palp 


Femur 


5-40 


3-44 


3-36 


4-28 


2-20 


Patella 


2-20 


1-48 


1-28 


1-36 


1-38 


Tibia 


5-68 


3-16 


3-08 


4-20 


1-08 


Metatarsus 


3-48 


2-56 


3-24 


4-58 






Tarsus 


1-22 


0-92 


1-04 


1-20 


1-88 



Total 17-98 11-56 12-00 15-62 6-54 

Ratios: AM : AL: : PM : PL :: 19 : 1 1 : 8 : 1 1; AL PM PL :: 10 : 13; AM : CL :: 19 : 5. 

VARIATION. Male total length varies from 6-24 to 7-44 mm, carapace length 2-72-3-36 mm (five 
specimens); female total length 6-0-6-96 mm, carapace length 2-56-3-36 mm (10 specimens). 

In two females the arthrodial membrane between the chelicerae and the clypeus is evident as a 
narrow white band, whereas it is broad and conspicuous in other specimens. However, to judge 
from the set of the chelicerae it is apparent that the phenomenon is the result of postmorten 
changes. 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 



115 




Fig. 6. Spartaeus wildtrackii sp. n., paratype cJ, A, palp, ventral view; B, palpal tibia, dorsal view; C, 
cheliceral teeth; D, palp, retrolateral view; E, chelicerae, maxillae and labium. Abbreviations: rta, 
retrolateral tibial apophysis, tf, tegular furrow; va, ventral apophysis. 



NATURAL HISTORY. S. wildtrackii has so far only been found in lowland rainforest on the trunks of 
large trees covered in lichen or moss, against which the spiders are well camouflaged. A number of 
specimens were seen both at night and during the day resting on bark beneath large silken webs ca. 
5-6 cm constructed of glossy translucent silk that was often torn (Fig. 1,2). Their prey is unknown 
except for that of one specimen which was seen feeding on newly emerged moths, that were resting 
and evidently drying their wings. The spiders were not seen to jump, but it was noted that they were 
fast runners (P. D. Hillyard, pers. comm.). 

DISTRIBUTION. West Malaysia. 

MATERIAL EXAMINED. West Malaysia, P. D. Hillyard: Pahang State, Taman Negara, nr. Kuala Tahan, 
lowland primary rain forest: on tree trunks under sheet webs, 9.iii.l984, holotype , BMNH. 1985.8.16.2, 
paratypes 3 ??,BMNH. 1985.8. 16.3-5; on tree trunks, iii. 1985, paratypes 4??, 4<JcJ, BMNH 1985.8.16.6-13. 
Negeri Sembilan State, Pasoh Forest Reserve, on tree trunks in lowland primary rain forest, iii.1985, 
paratypes 3 <J<J, 9 ??, BMNH, 1985.8.16.14-25. 

ETYMOLOGY. This species is named for the BBC television programme 'Wildtrack' which has done 
much to encourage children to care for the environment and take an interest in natural history. 



116 F. R. WANLESS 

REMARKS. 1 . Postepigynal furrows (Fig. 5D) are an unusual feature of salticid epigynes and to date 
have only been found in one other spartaeine i.e. Gelotia bimaculata Thorell. Their function is 
uncertain, but they may form part of the supporting mechanism that holds the male palp in 
position during copulation. 

2. Loerbroks ( 1 984) has recently drawn attention to conspicuous similarities in palpal structure 
between Misumena vatia (Clerck), a crab spider (Family Thomisidae) and Phaeacius Koch, a genus 
of flattened salticid that has also been classified in the Spartaeinae. Futhermore, he has shown that 
in M . vatia, and probably all other thomisids, the ventral apophysis locks into the regular ridge as 
the palpal elements expand and rotate during copulation. The ventral apophysis (Figs 6A; 13F) 
and tegular furrow (Fig. 6A) ( = tegular ridge of Loerbroks) characteristic of all spartaeines are 
evidently homologous with those of thomisids and probably function in a similar manner. In M. 
vatia the inner surface of the ventral apophysis is covered in papillae (see Loerbroks 1984, Fig. 6) 
that evidently reduce friction between the apophysis and the rotating tegulum. Similar papillae 
might therefore be expected on the ventral apophysis of 5. wildtrackii, but are absent (Fig. 18A). 
However, spicule-like papillae do occur on the inner surface of the retrolateral tibial apophysis 
(Fig. 18B-D) and presumbly they too could reduce surface friction, although on the otherhand 
they may serve to prevent the apophysis from sliding out of position during copulaton. 

Although genital structures are not generally used in assessing relationships at the familial level, 
they are considered here because ventral apophyses and tegular furrows are not known to occur in 
other spider families. They may have arisen independently, but as Loerbroks (1984) has already 
stated they may provide evidence of a phylogenetic link between salticids and thomisids. 
Additional evidence is provided by Homann (1971) who has shown that the anterior median eyes 
(principal eyes) of most spiders are small and have few visual cells, whereas those of salticids and 
thomisids are unique in possessing many visual cells which provide for 'sharp vision'. These optical 
similarities may be convergent as Homann regards thomisid eyes as being structurally closer to 
those of wolf spiders (Family Lycosidae). Future studies will have to take account of the genitalial 
similarities noted by Loerbroks since these can be interpreted as supportive of a sister group 
relationship between salticids and thomisids. 

Genus MINTONIA Wanless 
Mintonia Wanless, 1984a: 157. [definition, diagnosis and key to species]. 

This small oriental genus comprises nine species including two new taxa described below. The 
majority of species have been collected from Borneo, but the genus is also known from Java, 
Sumatra and Peninsular Malaya. Males are of particular interest because they possess femoral 
organs (Fig. 9D; 19A-C), a presumptive sex pheromone dispersal site, and retrolateral tibial 
apophyses, some of which bear openings (Fig. 9E arrowed; 20A, B). Unfortunately nothing is 
known of their natural history. 

Mintonia melinauensis Wanless 
(Fig. 7A-E) 

Mintonia melinauensis Wanless, 1984a: 165, $ holotype, Sarawak (BMNH) [examined]. 

DIAGNOSIS. Males can be recognized by the heavy inward curving embolus and by the form of the 
retrolateral tibial apophysis (see Wanless, 1984a, Fig. 13); females by the structure of the epigyne 
(Fig. 7E) which is clearly different from that of other females of the genus. 

FEMALE, formerly undescribed, in fair condition. Carapace (Fig. 7A, D): weakly iridescent under 
some angles of illumination; orange-brown lightly mottled black with paler lateral markings and 
band from fovea to posterior margin; rubbed, but otherwise clothed in whitish pubescent hairs. 
Eyes: laterals with black surrounds; fringed by whitish hairs. Clypeus: lightly tinged with some 
black; sparsely clothed in fine whitish hairs with several long bristles. Chelicerae: orange-brown 
suffused with some black proximally; shiny, thinly clothed in long pale orange hairs with modera- 
tely dense promarginal scopulae; promargin with three teeth, retromargin with six (Fig. 7B). 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 



117 




B 






Fig. 7. Mintonia melinauensis Wanless, 9: A, dorsal view; B, cheliceral teeth; C, sternum; D, carapace, 

lateral view; E, epigyne. 

Maxillae: light orange-brown with whitish yellow inner distal margins. Labium: brownish orange 
tipped whitish yellow. Sternum (Fig. 7C): pale yellow with darker margins; thinly clothed in fine 
hairs. Coxae: pale greyish yellow. Abdomen: rubbed; whitish yellow with vague greyish markings; 
spinnerets whitish yellow with outer sides of anteriors tinged black. Legs: moderately long and 
robust; whitish yellow to orange brown; spines strong and numerous. Spination of leg I; metatar- 
sus v 2-2-1, p 1-0-1, d 0-0-2, r 1-0-0; tibia v 2-2-2, p 1-0-1, r 0-0-1; patella p 0-1-0, r 0-1-0; 
femur d 0-2-3. Palp: whitish yellow with sooty markings except for pale orange brown tarsi. 
Epigyne (Fig. 7E): clothed in fine pale yellowish hairs. 

Dimensions (mm): total length 5-2; carapace length 2-24, breadth 1-76, height 1-44; abdomen 
length 3-04; eyes, anterior row 1-64, middle row 1-48, posterior row 1-64; quadrangle length 1-2 
(53% of carapace length). 



Leg 


1 


2 


3 


4 


Palp 


Femur 


1-48 


1-46 


1-48 


1-80 


0-80 


Patella 


0-86 


0-81 


0-71 


0-76 


0-48 


Tibia 


1-12 


1-04 


1-12 


1-44 


0-52 


Metatarsus 


0-96 


0-94 


1-16 


1-60 






Tarsus 


0-58 


0-57 


0-60 


0-68 


0-72 



Total 



5-00 



4-82 



5-07 6-28 



2-52 



Ratios: AM : AL : PM : PL:: 13:8:5:8; AL PM PL :: 7-5-8-5; AM : CL :: 13 : 2-5. 
DISTRIBUTION. Sarawak. 



118 



F. R. WANLESS 





rta 





Fig. 8. Mintonia caliginosa sp. n., holotype $: A, dorsal view; B, carapace, lateral view; C, palp, ventral 
view; D, palp, retrolateral view; E, palp, dorsal view. Abbreviations: e, embolus; rta, retrolateral tibial 
apophysis; v, vacuole; va, ventral apophysis. 

MATERIAL EXAMINED. Type data given in synonymy. Sarawak: Gunung Mulu National Park, Environs of 
camp 3, moss forest, 1$, from moss covered tree trunk, 27.5.78, F. R. Wanless, Royal Geographic Society/ 
Sarawak Government Expedition. (BMNH). 

Mintonia caliginosa sp. n. 

(Fig. 8A-E;19A-C) 

DIAGNOSIS. A distinctive species easily recognized by the long retrolateral tibial apophysis 
(Fig. 8D). 

FEMALE. Unknown. 

MALE HOLOTYPE, rubbed otherwise in fair condition. Carapace (Fig. 8A, B): orange-brown lightly 
tinged and mottled black; shiny and weakly iridescent under some angles of illumination. Eyes: 
laterals with black surrounds; fringed by whitish and pale amber hairs, mostly rubbed. Clypeus 
edged black below anterior median eyes; rubbed a few whitish hairs remaining. Chelicerae: light 
brown with black markings, shiny, thinly clothed in scattered fine hairs; promargin with three 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 1 1 9 

teeth, retromargin with four or five. Maxillae: light yellowish brown. Labium: blackish edged pale 
yellow-brown. Sternum: yellow-brown, shiny, thinly clothed in fine greyish hairs. Abdomen: 
yellow-brown lightly tinged and mottled black with a poorly defined orange-brown scutum and 
chevrons dorsally, and three rather vague longitudinal bands ventrally; spinnerets yellow-brown 
lightly tinged black. Legs: moderately long and slender; femoral organ (Fig. 19A-C) a low 
tubercle; yellow-brown tinged with some black except metatarsi and tarsi which are darker 
orange-brown tinged black, also on underside of femora I-II a transverse blackish patch; spines 
strong and numerous. Spination of leg I: metatarsus v 2-0-0, p 1-1-1, d 0-1-2, r 1-1-1; tibia v 
2-2-2, p 1-1-0, d 1-1-0, r 1-1-0; patella p 0-1-0, r 0-1-0; femur d 0-2^. Palp (Fig. 8C-E): the 
retrolateral tibial apophysis is broken at point arrowed in Fig. 8E; however, note that in Fig. 8C i.e. 
the same palp but drawn from a different angle, the appearance of the retrolateral tibial apophysis 
has been reconstructed from the apophysis of the other palp. 

Dimensions (mm): total length 3-8; carapace length 1-76, breadth 1-36, height 1-1; abdomen 
length 1-92; eyes, anterior row 1-27, middle row 1-2, posterior row 1-35; quadrangle length 0-94 
(53% of carapace length). 



Leg 


1 


2 


3 


4 


Palp 


Femur 


1-16 


1-12 


1-19 


1-46 


0-60 


Patella 


0-60 


0-60 


0-52 


0-56 


0-32 


Tibia 


0-84 


0-80 


0-88 


1-16 


0-24 


\^of Q t Q *C1 1 C 


0-80 


0-78 


0-88 


1 -78 




IVItldltll oUS 

Tarsus 


\J OU 

0-52 


u / o 

0-48 


u oo 

0-56 


i _ <> 

0-60 


0-72 



Total 3-90 3-78 4-03 5-06 1-88 

Ratios: AM : AL : PM : PL :: 9-5 : 5-5 : 3-5 : 5-5; AL PM PL :: 6-7; AM : CL :: 9-5 : 3-3. 
DISTRIBUTION. Borneo, Sabah. 

MATERIAL EXAMINED. Borneo: Sabah, Tuaran Division, Mt. Kinabalu National Park, Power Station 
Layang Layang, cloud forest, holotype <$, 2000-2800 m, 7.H.1976 P. T. Lehtinen, (TU, Turku). 

REMARK. The presence of a vacuole in the base of the retrolateral tibial apophysis suggests that 
there is probably a distal opening. There is insufficient material for this to be confirmed by SEM. 

ETYMOLOGY. The specific name is from the Latin meaning misty, cloudy places. 



Mintonia silvicola sp. n. 
(Fig. 9A-G) 

DIAGNOSIS. M. silvicola seems to be most closely related to M. tauricornis Wanless, but may be 
readily distinguished by the syringe-shaped retrolateral tibial apophysis (Fig. 9E). 

FEMALE. Unknown. 

MALE HOLOTYPE, rubbed, also right leg I missing, otherwise in fair condition. Carapace (Fig. 9A, 
B): weakly iridescent under some angles of illumination; orange-brown with faint blackish 
mottling on sides. Eyes: laterals with black surrounds; anteriors fringed by whitish hairs. Clypeus: 
orange-brown with blackish margin below anterior median eyes and vague yellow-brown mark- 
ings clothed in whitish hairs below anterior laterals. Chelicerae: yellow-brown, shiny, clothed in 
white hairs proximally and scattered brown hairs distally with dense promarginal scopulae; 
promargin with three teeth, retromargin with eight (Fig. 9G). Maxillae: yellow-brown with inner 
distal margins paler. Labium: yellow-brown faintly tinged grey. Sternum: pale yellow with vague 
darker margins; thinly clothed in fine pale yellow hairs. Coxaei pale yellow. Abdomen: pale yellow 



120 



F. R. WANLESS 




B 



Fig. 9. Mintonia silvicola sp. n., holotype <$: A, dorsal view; B, carapace, lateral view; C, palp, ventro- 
lateral view; D, leg I; E, palp, retrolateral view; F, palp, ventral view; G, cheliceral teeth. Abbreviation: 
fo, femoral organ. 



with vague sooty markings and two pairs of sigilla; rubbed; spinnerets moderately long, yellow- 
brown. Legs: moderately long and slender; specialized prey capture and metatarsal setae lacking; 
femoral organ appearing as a low dark amber mound; legs I pale yellow to yellow-brown with 
underside of femora tinged black; other legs pale yellow grading to yellow-brown distally with 
ventral longitudinal grey stripe on tibiae II-IH; spines numerous and moderately strong. Spination 
of leg I: metatarsus v 2-0-0, r 1-1-1, d 0-2-2, p 1-1-1; tibia v 2-2-2, p 1-1-0, d 1-1-0, r 1-1-0; 
patella p 0-1-0, r 0-1-0; femur d 0-2^. Palp (Fig. 9C, E, F): element M2 lies above the embolus 
and the tegular ledge is poorly developed; the opening of the retrolateral tibial apophysis (arrow, 
Fig. 9E) is distinct. 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 1 2 1 

Dimensions (mm): total length 5-1; carapace length 2-24, breadth 1-88, height 1-44; abdomen 
length 2-6; eyes, anterior row 1-71, middle row 1-48, posterior row 1-64; quadrangle length 1-32 
(58% of carapace length). 



Leg 


1 


2 


3 


4 


Palp 


Femur 


1-72 


1-72 


1-72 


2-04 


0-84 


Patella 


0-88 


0-84 


0-76 


0-80 


0-40 


Tibia 


1-36 


1-28 


1-34 


1-64 


0-34 


\^f*t d t d rcnc 


1 -78 


1 -96 


1 -44 


1 -80 




IVlCLd. Ldl S US 

Tarsus 


i 'i 

0-64 


1 O 

0-64 


1 HH 

0-72 


I OU 

0-76 


1-12 



Total 5-88 5-74 5-98 7-04 2-70 

Ratios: AM : AL : PM : PL :: 14 : 8 : 5-4 : 8; AL PM PL :: 8-9-5; AM : CL :: 14 : 3. 
DISTRIBUTION. West Malaysia. 

MATERIAL EXAMINED. West Malaysia: Pahang State, Taman Negara, holotype $, from buttress of large tree, 
lowland rain forest nr. Kuala Tahan, 3-10.iii.1984. P. D. Hillyard, BMNH. 1985.8.21.1. 

ETYMOLOGY. The specific name is from the Latin meaning inhabiting woods. 

Genus TARAXELLA Wanless 
Taraxella Wanless, 1984a: 155. [definition and diagnosis]. 

This genus was originally erected on the basis of a single male of Taraxella solitaria Wanless, 
from Sarawak. Subsequent collections have produced four new species, described below, that 
necessitate modifications to the original generic definition. 

DEFINITION. Spiders small to medium in size, i.e. between 2-0 and 8-0 mm in length; males some- 
times with conspicuous encircling band on the carapace; sexual dimorphism sometimes evident in 
colour patterns. 

Carapace, high, longer than broad, widest at about level of coxae II-III; fovea long and sulci- 
form, apex at level of centre of posterior lateral eyes. Eyes: anterior medians more or less level or 
weakly procurved in frontal view; posterior medians small to relatively large; posterior laterals 
with outer margins of lenses set inside or at level of, lateral margins of carapace when viewed from 
above; entire quadrangle length between 57-65% of carapace length. Clypeus: low to moderately 
high. Chelicerae: promargin with five or seven teeth, retromargin with seven or nine denticles. 
Legs: moderately long and slender; femoral organs lacking; specialized prey capture tarsal setae 
and filamentous metatarsal setae also lacking. Female palps: moderately long and slender with 
apical claw. Epigynes: interspecifically distinct, see descriptions; vulvae not examined, insufficient 
material. Male palps: complex and interspecifically distinct; retrolateral tibial apophyses complex, 
sometimes bifid with sharp slender prongs, or evidently reduced with associated stout setae; 
apophyses X and Y variable in development; embolus short slender and gently curved, and for the 
most part obscured, in ventral view, by tegular apophyses X and occasionally Y; tegular furrow 
and ventral apophysis usually conspicuous; Ml, see Wanless 19840, a delicate fan-shaped lamella 
that protrudes beyond the distal edge of the tegulum. Expanded palps not examined. 

DIAGNOSIS. Distinguished from other spartaeines by the conformation of the embolus of the 
male palp which is almost completely obscured, in ventral view, by tegular apophyses 'X' and 
occasionally 'Y'. 

An identification key is not provided as the five known species are easily separated from one 
another by the structure of the palpal organs and epigynes. 

INTERSPECIFIC RELATIONSHIPS. To judge from the structure of the tibia of the male palpal organs T. 
solitaria, T. petrensis sp. n., and T. hilly ardi sp. n., form a closely related group since they all possess 
retrolateral tibial apophyses with a sharp dorsal prong. T. sumatrana sp. n., and T. reinholdae 



122 



F. R. WANLESS 



am 



al 




B 



Fig. 10. Taraxella hillyardi sp. n., holotype c?: A, dorsal view; B, carapace, lateral view; C, leg I; D, 
palpal tibia, dorsal view; E, palp, retrolateral view; F, palp, ventral view. Abbreviations: al, anterior 
lateral eye; am, anterior median eye; dh, distal haematodocha; e, embolus; pi, posterior lateral eye; pm, 
posterior median eye; tf, tegular furrow; va, ventral apophysis. 

would also appear to form a natural group as they possess conspicuous fringes of unusually stout 
setae, and lack retrolateral tibial apophyses with sharp dorsal prongs. 



Taraxella hillyardi sp. n. 
(Fig. 10 A, F) 

DIAGNOSIS. T. hillyardi seems to be most closely related to T. petrensis and T. solitaria, but can be 
easily distinguished by the bifid retrolateral tibial apophysis (Fig. 10E). 

FEMALE. Unknown. 

MALE HOLOTYPE, in fair condition. Carapace (Fig. 10A, B): weakly iridescent under some angles of 
illumination; light orange-brown tinged black with a broad yellowish encircling band on sides and 
also a black marginal band extending posteriorly from level of coxae I; rubbed. Eyes: laterals with 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 1 23 

black surrounds; sparsely fringed in pale amber and whitish hairs. Clypeus: greyish with black 
markings below anterior median eyes and whitish yellow stripes below anterior laterals; sparsely 
fringed by whitish hairs. Chelicerae: yellow-brown with extensive patches on facies; shiny; thinly 
clothed in fine clear hairs and some blackish ones with dense promarginal scopulae; promargin 
with five teeth; retromargin with eight denticles. Maxillae andlabium: pale yellow with vague sooty 
markings. Coxae: yellow-brown. Abdomen: dorsum and sides pale yellow-brown suffused and 
mottled black, venter pale yellow brown suffused black in region of tracheal spiracle; spinnerets 
moderately long; anteriors and posteriors suffused black, medians pale yellow. Legs Fig. IOC): 
moderately long and slender; generally yellow-brown tinged with some black, with incomplete 
annuli on femora and blackish tibiae particularly of legs I and IV; spines moderately strong and 
numerous. Spination of leg I: metatarsus v 2-0-0, p. 1-1-1 , d 0-2-2, r 1-1-1 ; tibia v 2-2-2, p 0-1-1 , 
d i_i_o, r 1-1-0; patella p 0-1-1, r 0-1-0; femur d 0-2-3, p 0-0-1. Palp (Fig. 10D-F): yellow- 
brown to orange-brown mottled black with patches of white hairs on patella and apices of femur, 
otherwise clothed in black hairs and scattered white ones with greyish scopula on cymbium. 

Dimensions (mm): total length 3-9; carapace length 1-76, breadth 1-53, height 1.2; abdomen 
length 1-84; eyes, anterior row 1-56, middle row 1-24, posterior row 1-52; quadrangle length 1-07 
(60% of carapace length). 



Leg 


1 


2 


3 


4 


Palp 


Femur 


1-36 


1-28 


1-28 


1-64 


0-68 


Patella 


0-76 


0-63 


0-56 


0-64 


0-33 


Tibia 


1-12 


0-96 


1-00 


1-39 


0-36 


Metatarsus 


1-02 


0-96 


1-12 


1-60 






Tarsus 


0-53 


0-52 


0-56 


0-66 


0-72 



Total 4-79 4-35 4-52 5-93 2-09 

Ratios: AM : AL : PM : PL :: 13 : 7 : 3 : 7; AL PM PL :: 7-8; AM : CL :: 13 : 3. 
DISTRIBUTION. West Malaysia. 

MATERIAL EXAMINED. West Malaysia: Gunong Jerai, (Kedeh), ca. 700 m, holotype <$, shrub layer, along forest 
edge during middle of dry season, 15.ii.1983, P. D. Hillyard(BMNH. 1985.9.5.2). 

ETYMOLOGY. This species is named after my colleague Mr P. D. Hillyard, BMNH, who collected 
many of the new species described in this paper. 

Taraxella petrensis sp. n. 
(Fig. 11A-J) 

DIAGNOSIS. T. petrensis seems to be most closely related to T. solitarius and T. hillyardi sp. n., but 
may be easily separated by the broad flange of the retrolateral tibial apophysis (Fig. 1 U) in males 
and the structure of the epigyne (Fig. 1 ID) in females. 

MALE HOLOTYPE, rubbed otherwise in good condition. Carapace (Fig. 1 1 A, F): shiny and weakly 
iridescent in eye region; dark orange-brown suffused black with broad encircling creamy white 
band. Eyes: laterals with black surrounds; anteriors sparsely fringed in greyish hairs. Clypeus: 
creamy white with black spots near lower rims of anterior median eyes. Chelicerae: shiny black 
except for orange-brown inner margins; sparsely clothed in greyish hairs with dense promarginal 
scopulae; promargin with five teeth, retromargin with nine denticles (Fig. 11C). Maxillae and 
labium: greyish yellow faintly tinged black. Sternum: pale yellow-brown with darker margins; 
thinly clothed in light brownish hairs. Coxae: yellow-brown tinged grey. Abdomen: yellow-brown 
with dorsum and sides mottled black, also a vague black patch in area of tracheal slit; rubbed; 
spinnerets moderately long, black except for light greyish medians. Legs: moderately long and 
slender; yellow-brown heavily suffused black, especially on femora, patellae and tibiae; shiny and 



124 



F. R. WANLESS 




Fig. 11. Taraxella petrensis sp. n., holotype (J: A, dorsal view; C, cheliceral teeth; F, carapace, lateral 
view; G, palpal tibia, dorsal view; H, palp, ventral view; J, palp, retrolateral view. Paratype $: 
B, cheliceral teeth; E, carapace, dorsal view; I, leg I. Abbreviation: e, embolus; 'x' and 'y' tegular 
apophyses. 

iridescent under some angles or illumination; spines strong and numerous. Spination of leg I: 
metatarsus v 2-2-2, p 1-0-0, d 0-1-2, r 1-0-0; tibia v 1-3-2, p 0-1-1, d 1-1-0, r 0-0-1; patella p 
0-1-0, r 0-1-0; femur d 0-2-3. Palp (Fig. 1 1G, H, J): yellow to orange-brown suffused with some 
black especially on femur and cymbium; clothed in pale grey and black hairs. 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 1 25 

Dimensions (mm): total length 3-72; carapace length 1-76, breadth 1-56, height 1-24; abdomen 
length 2-16; eyes, anterior row 1-6; middle row 1-21, posterior row 1-52; quadrangle length 1-08 
(61 % of carapace length). 



Leg 


1 


2 


3 


4 


Palp 


Femur 


1-52 


1-34 


1-36 


1-72 


0-76 


Patella 


0-76 


0-60 


0-60 


0-66 


0-32 


Tibia 


1-24 


1-00 


1-04 


1-42 


0-36 


\ji t 


1-19 


1 -09 


1 1 X 


1 -fis 




ivieiaiarsus 
Tarsus 


1 1Z 

0-60 


1 \JZ. 

0-56 


1 1 

0-56 


1 U J 

0-70 


0-84 



Total 5-24 4-52 4-74 6-15 2-28 

Ratios: AM : AL : PM : PL :: 13-5 : 7-5 : 2-5 : 7-5; AL PM PL :: 7-5 : 7; AM : CL :: 13-5 : 5. 

FEMALE PARATYPE, in fair condition. General habitus as in male except encircling cephalic band 
lacking. Carapace (Fig. 1 IE): light orange-brown lightly and finely reticulated black in eye region 
with blackish mottling on sides, also weakly iridescent under some angles of illumination; rubbed. 
Eyes: generally as in male, but sparsely fringed by pale amber hairs. Clypeus: light orange-brown 
faintly reticulated black; bald except for scattered fine marginal hairs and several long stiff hairs 
including usual triad in lower space between anterior median eyes. Chelicerae: yellow-brown 
lightly tinged with some black; shiny; sparsely clothed in brown hairs with dense promarginal 
scopulae; promargin with five teeth, retromargin with nine denticles (Fig. 11B). Maxillae and 
labium: yellow-brown. Sternum: pale yellow-brown with darker margins; sparsely clothed in fine 
hairs centrally and darker, longer ones towards margins. Abdomen: generally as in male except 
venter pale yellow-brown with scattered dark brown simple hairs and vague light greyish feathery 
hairs, otherwise rubbed; spinnerets similar to male, yellow-brown tinged black. Legs (Fig. Ill): 
moderately long and slender; light orange-brown faintly tinged with some black; sparsely clothed 
in brownish hairs; spines strong and numerous. Spination of leg I: metatarsus v 2-2-0, p 1-0-1, d 
0-1-2, r 0-0-1; tibia v 2-3-1, p 0-1-1; patella p 0-1-0; femur d 0-2-2. Epigyne (Fig. 1 ID). 

Dimensions (mm): total length 4-32; carapace length 1-96, breadth 1/72, height 1-32; abdomen 
length 2-28; eyes, anterior row 1-76, middle row 1-36, posterior row 1-72; quadrangle length 1-2 
(61% of carapace length). 



Leg 


1 


2 


3 


4 


Palp 


Femur 


1-58 


1-44 


1-44 


1-76 


0-76 


Patella 


0-84 


0-74 


0-64 


0-70 


0-43 


Tibia 


1-18 


1-00 


1-08 


1-46 


0-48 


Metatarsus 


1-04 


1-00 


1-20 


1-68 






Tarsus 


0-60 


0-56 


0-63 


0-64 


0-67 



Total 5-24 4-74 4-99 6-24 2-34 

Ratios: AM : AL : PM : PL :: 15 : 8 : 2 : 8; AL PM PL :: 8-9; AM : CL :: 15 : 2. 
DISTRIBUTION. West Sumatra. 

MATERIAL EXAMINED. West Sumatra: Harau Nature Reserve, near Payakumbu, holotype c?, on rock walls on 
edge of forest, ca. 600 m, ii. 1985, P. D. Hillyard, (BMNH. 1985.9.5. 1); Taram, near Payakumbu, in secondary 
forest litter, paratype ?, ii.1985, P. D. Hillyard, (BMNH. 1985.9.5.2). 

ETYMOLOGY. The specific name is from the Latin meaning rocky places. 

Taraxella sumatrana sp. n. 

(Fig. 12A-J) 

DIAGNOSIS. Easily separated from other species of Taraxella by the dorsal fringe of stout setae on 
the palpal tibia (Fig. 12G, J) in males and by the structure of the epigyne in females (Fig. 12D). 



126 



F. R. WANLESS 




Fig. 12 Taraxella sumatrana sp. n., holotype ?: A, dorsal view. B, cheliceral teeth; D, epigyne; F, 
sternum; I, maxillae and labium. Paratype <$: C, cheliceral teeth; E, carapace, lateral view; G, palpal 
tibia, dorsal view; H, palp, ventral view; J, palp, retrolateral view. Abbreviations: e, embolus; Ml, 
fan-shaped element of distal haematodocha. 

FEMALE HOLOTYPE, in fair condition. Carapace (Fig. 12A): yellow-brown faintly tinged and 
mottled black with a metallic sheen under some angles of illumination; rubbed except for some 
scattered brown hairs. Eyes: laterals with black surrounds; fringed by light brownish hairs and 
some whitish ones below anterior median eyes. Clypeus yellow-brown tinged black with a few 
black bristles. Chelicerae: pale yellow-brown, shiny, sparsely clothed in brown-black hairs and 
some bristles with dense promarginal scopulae; promargin with six teeth, retromargin with eight 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 1 27 

denticles (Fig. 12B). Maxillae and labium: pale yellow-brown. Sternum (Fig. 12F): pale yellow- 
brown with darker margins; thinly clothed in stiff brown hairs. Coxae: pale yellow-brown. 
Abdomen: pale yellow-brown tinged and mottled black with vague chevrons dorsally; venter pale 
greyish yellow; mostly rubbed otherwise sparsely clothed in patches of recumbent dull amber 
lanceolate hairs, thinly interspersed with erect black hairs; spinnerets moderately long, pale yellow 
brown except anteriors tinged with some black. Legs: moderately long and slender; generally pale 
yellow-brown faintly tinged with some black; thinly clothed in pale greyish and black hairs; spines 
moderately strong and numerous. Spination of leg I: metatarsus v 2-0-0, r 0-1-1 , d 0-1-2, p 1-1-1 ; 
tibia v 1-3-2, p 0-1-1; patella p 0-1-0; femur d 0-2-3. Epigyne (Fig. 12D): clothed in dark grey 
hairs. 

Dimensions (mm): total length 4-44; carapace length 2-0, breadth 1-68, height 1-32; abdomen 
length 2-32; eyes, anterior row 1-72, middle row 1-38, posterior row 1-61; quadrangle length 1-12 
(56% of carapace length). 



Leg 


1 


2 


3 


4 


Palp 


Femur 


1-52 


1-36 


1-36 


1-76 


0-76 


Patella 


0-78 


0-68 


0-62 


0-68 


0-40 


Tibia 


1-12 


0-96 


1-00 


1-40 


0-52 


IMpf H tfl TQ11Q 


1-02 


0-96 


1.1 1 


1 % 




1 VI I/ Id Id 1 T LI .> 

Tarsus 


1 \ t ~ 

0-56 


\J .s\J 

0-56 


1 11 

0-58 


1 J\J 

0-70 


0-66 



Total 5-00 4-52 4-67 6-10 2-34 

Ratios: AM : AL : PM : PL :: 14 : 8 : 4 : 8; AL PM PL :: 7-7-5; AM : CL :: 14 : 3. 

MALE PARATYPE, abdomen and legs IV missing, otherwise in fair condition. Similar to female 
except for the following. Carapace (Fig. 12E): heavily suffused and mottled black especially on 
sides; rubbed except for shining violet hairs behind anterior eyes. Clypeus: with scattered fine 
whitish hairs. Chelicerae: yellow-brown with sooty markings, shiny, clothed in scattered black 
hairs with dense promarginal scopulae; promargin with five teeth retromargin with seven denticles 
(Fig. 12C). Spination of leg I: metatarsus v 2-2-2, r 1-0-0, d 0-2-2, p 1-0-0; tibia v 1-3-2, p 0-1-1, 
r 1-0-1, d 1-0-0; patella p 0-1-0, r 0-1-0; femur d 0-2-3. Palp (Fig. 12G, H, J): femur and patella 
pale yellowish lightly suffused black; tibia and cymbium light to dark amber mottled with some 
black; clothed in brownish hairs with patch of white hairs on patella and apices of femora. 

Dimensions (mm): total length ?; carapace length 1 -92, breadth 1 -22, height 1 -32; abdomen length 
?; eyes, anterior row 1-64, middle row 1-29, posterior row 1-52; quadrangle length 1-12 (58% of 
carapace length). 



Leg 
Femur 
Patella 
Tibia 
Metatarsus 
Tarsus 


1 
1-58 
0-76 
1-26 
1-16 
0-62 


2 
1-44 
0-68 
1-04 
1-08 
0-60 


3 
1-40 
0-60 
1-08 
1-22 
0-64 


4 Palp 
fl-RO 


0-^fi 


0-d.fi 




0-92 





Total 5-38 4-84 4-94 2-54 

Ratios: AM : AL : PM : PL :: 14 : 8 : 4 : 8; AL PM PL :: 6-7; AM : CL :: 14 : 5. 
DISTRIBUTION. Sumatra. 

MATERIAL EXAMINED. Sumatra, Bohorok, Gunung Leuser Reserve: holotype ?, in litter, 14.vi. 1983, P. R. 
Deeleman and C. L. Deeleman- Reinhold; paratype , in bamboo litter on plateau with large bamboo trees, 
50 m, lO.ii. 1983, P. R. Deeleman and S. Djojosudharma (Rijksmuseum van Natuurlijke Histoire, Leiden). 

ETYMOLOGY. The specific name refers to the country in which the holotype was collected. 



128 



F. R. WANLESS 




B 



Fig. 13. Taraxella reinholdae sp. n., holotype $: A, dorsal view; B, carapace, lateral view; C, leg I; D, 
palpal tibia, dorsal view; E, cheliceral teeth; F, palp, ventral view; G, palp, retrolateral view. Abbrevi- 
ations: M 1 , fan-shaped element of distal haematodocha; tf, tegular furrow; va, ventral apophysis; 'x' 
and 'y', tegular apophyses. 



Taraxella reinholdae sp. n. 
(Fig. 13A-G) 

DIAGNOSIS. Easily separated from other species of Taraxella by the conspicuous tuft of stout setae 
arising from the retrolateral surface of the palpal tibial apophysis (Fig. 13D, F, G). 

FEMALE. Unknown. 

MALE HOLOTYPE in fair condition. Carapace (Fig. 13 A, B): mottled black with metallic sheen and 
pale yellowish green markings on thoracic part; irregularly clothed, in scattered black and white 
hairs with a central white haired stripe behind anterior median eyes. Eyes: laterals with black 
surrounds; fringed by white hairs with scattered dark grey hairs around laterals and upper rims of 
anterior medians. Clypeus: clothed in scattered white hairs. Chelicerae: black with metallic sheen 
except for light greyish amber inner margins; sparsely clothed in light greyish hairs with moderately 
dense promarginal scopulae; promargin with five teeth, retromargin with seven denticles (Fig. 
13E). Maxillae and labium: pale yellow-brown heavily mottled black with whitish inner distal 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 1 29 

margins on maxillae and labial tip. Sternum: yellow-brown suffused black; shiny, with scattered 
fine blackish hairs. Coxae: pale greyish yellow suffused with some black. Abdomen: dorsum and 
sides yellowish green heavily tinged black; clothed in brown-black hairs with vague patches of 
whitish hairs laterally; venter pale yellow with black surrounds, clothed in grey black hairs; 
spinnerets black with blackish hairs. Legs (Fig. 1 3C): moderately long and slender; generally black 
except for pale yellowish brown tarsi and yellow-brown annuli on metatarsi, tibiae and patellae; 
spines strong and numerous. Spination of leg I: metatarsus v 2-0-0, p 1-0-2, d 1-1-2; tibia v 2-2-2, 
p 1-0-1 , d 1-1-0; patella p 0-1-0, r 0-1-0; femur d 0-2-3. Palp (Fig. 1 3D, F, G): patella pale yellow 
tinged with some black, other segments heavily mottled black; clothed in black hairs with patch of 
whitish hairs on patella and apices of femora. 

Dimensions (mm): total length ca. 3-2 (bent); carapace length 1-52, breadth 1-44, height 1-16; 
abdomen length 1-68; eyes, anterior row 1-48, middle row 1-16, posterior row 1-40; quadrangle 
length 1-0 (65% of carapace length). 



Leg 

Femur 
Patella 
Tibia 
Metatarsus 
Tarsus 


1 
1-28 
0-66 
1-02 
0-96 
0-52 


2 
1-18 
0-58 
0-84 
0-88 
0-48 


3 
1-20 
0-54 
0-88 
0-96 
0-52 


4 
1-56 
0-60 
1-20 
1-42 
0-61 


Palp 
0-64 
0-32 
0-32 


0-72 



Total 4-44 3-96 4-10 5-39 2-00 

Ratios: AM : AL : PM : PL :: 7 : 12-5 : 2 : 7; AL PM PL :: 8-7; AM : CL :: 12-5 : 3-5. 
DISTRIBUTION. Borneo, Sarawak. 

MATERIAL EXAMINED. Sarawak: Bako National Park, holotype $, in litter, in swampy forest, 29-30.iii. 1 985. C. 
L. Deeleman-Reinhold and P. R. Deeleman, (RNH, Leiden). 

ETYMOLOGY. This species is named for one of the collectors Dr C. L. Deeleman-Reinhold. 



Acknowledgements 

I would like to thank Dr C. L. Deeleman-Reinhold, Ossendrecht, Holland and Dr P. Lehtinen, University of 
Turku, Turku, Finland for allowing me to study their collections of oriental salticids. 

I am also indebted to my colleague Mr P. D. Hillyard for collecting spiders in Indonesia and Malaysia 
and for providing photographs for Figs 1-2. Mrs F. Murphy, London kindly supplied the photograph for 
Figure 3. 

Finally I would like to thank Dr R. R. Jackson, University of Canterbury, Christchurch, New Zealand and 
Mr D. Macfarlane, Commonwealth Institute for Entomology, London for reading the manuscript. 



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webs, predatory versatility, and intraspecific interactions. N.Z. Jl. Zool. 

& (in press b). Predatory versatility and intraspecific interactions of spartaeine jumping spiders 



(Araneae: Salticidae): Brettus adonis, B. cingulatus, Cyrba algerina, and Phaeacius sp. n. N.Z. Jl. Zool. 
Legendre, R. & Llinares, D. 1970. L'accouplement de 1'araignee salticide Cyrba algerina (Lucas, 1846). Annls 

Soc. Hon. Hist. nat. Herault 110(4): 169-174. 
Loerbroks, A. L. 1984. Mechanik der Kopulationsorgane von Misumena vatia (Clerck, 1757) (Arachnida: 

araneae: Thomisidae). Verh. naturwisc. Ver. Hamburg (NF) 27: 383-403. 
Platnick, N. I. & Shadab, M. U. 1975. A revision of the spider genus Gnaphosa (Araneae: Gnaphosidae) in 

America. Bull. Am. Mus. nat. Hist. 155: 3-66. 
Wanless, F. R. 1978a. A revision of the spider genera Belippo and Myrmarachne (Araneae: Salticidae) in the 

Ethiopian Region. Bull. Br. Mus. nat. Hist. (Zool.) 33(1): 139 pp. 
19786. A revision of the spider genus Portia (Araneae: Salticidae). Bull. Br. Mus. nat. Hist. (Zool.) 34(3): 

83-124. 
1984a. A review of the spider subfamily Spartaeinae nom. n. (Araneae: Salticidae) with descriptions of 

six new genera. Bull. Br. Mus. nat. Hist. (Zool.) 46(2): 135-205. 

19846. A revision of the spider genus Cyrba (Araneae: Salticidae) with the description of a new presump- 
tive pheromone dispersing organ. Bull. Br. Mus. nat. Hist. (Zool.) 47(7): 445-481. 
1985. A revision of the spider genera Holcolaetis and Sonoita (Araneae: Salticidae). Bull. Br. Mus. nat. 

Hist. (Zool.) 48(4): 249-278. 

Manuscript accepted for publication 21 February 1986 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 



131 




Fig. 14. Spartaeus wildtrackii sp. n., paratype (J: femoral organ: A, x 316; B, x 820; C, x 1040. Scale 

bars urn. Arrow indicates direction of coxa. 



132 



F. R. WANLESS 







Fig. 15. Spartaeus wildtrackii sp. n., cJ, tibia II: A, dorsal view of distal region showing distribution of 
mytiliform organs and muscle attachment sites, x 570; B, mytiliform organs and muscle attachment 
sites, x 1600; C, detail of mytiliform organ, x 4200. Abbreviations: h, hair sockets; hi, hinge line; m, 
muscle attachment site; my, mytiliform organ. Scale bars urn. 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 



133 




Fig. 16. Spartaeus wildtrackii sp. n., J: A, femora I, mytiliform organs and seta, x 1000; B, femora II, 
detail of mytiliform organ, x 6200; C, metatarsus II, base of filimentous seta showing dendritic pore, 
x 10300; D, femora I, feathery setae, x 620. Scale bars ^m. 



134 



F. R. WANLESS 




Fig. 17. Spartaeus wildtrackii, sp. n., $: A, basal half of metatarsi II, lateral view showing fringe of 
filamentous setae; C, two filamentous setae, x 1300; D, E, F, mid region, distal region and tip of 
filamentous setae, x 7000, x 8000, x 8000; B, basal half of tarsi II, ventral view showing row of prey 
capture setae, arrowed; x 210; G, detail of prey capture setae, x 2000. Scale bars urn. 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 



135 




Fig. 18. Spartaeus wildtrackii sp. n., tibia of male palpal organ: A, ventral apophysis inner view, x 1000; 
B, retrolateral apophysis, outer view, x 100; C, retrolateral apophysis, inner view, x 100; D, detail of 
dorsal prong of retrolateral apophysis, x 350. Scale bars ^im. 



136 



F. R. WANLESS 




Fig. 19. Mintonia caliginosa sp. n., $: A, underside of femora I showing position of femoral organ, 
arrowed, x 75; B, femoral organ, x 550; C, femoral organ showing pores, x 2800. Scale bars A, mm; 
B, C, urn. 



NOTES ON SPIDERS OF THE FAMILY SALTICIDAE 



137 




Fig. 20. (A, B) Mintonia tauricornis Wanless, <$, palpal organ: A, tibia showing ventral apophysis 
and pore bearing retrolateral apophysis, x 150; B, tip of retrolateral apophysis showing pore, x 850. 
C, Portia labiata Thorell, <$, showing mytiliform field on dorsal surface of abdomen, x 730. 
Abbreviations: m, mytiliform organs; va, ventral apophysis; rta, retrolateral tibial apophysis. Scale 
bars |im. 



British Museum (Natural History) 

The birds of Mount Nimba, Liberia 



Peter R. Colston & Kai Curry-Lindahl 



For evolution and speciation of animals Mount Nimba in Liberia, Guinea and the Ivory Coast is 
a key area in Africa representing for biologists what the Abu Simbel site in Egypt signified for 
archaeologists. No less than about 200 species of animals are endemic to Mount Nimba. Yet, this 
mountain massif, entirely located within the rain-forest biome, is rapidly being destroyed by 
human exploitation. 

This book is the first major work on the birds of Mount Nimba and surrounding lowland 
rain-forests. During 20 years (1962-1982) of research at the Nimba Research Laboratory in 
Grassfield (Liberia), located at the foot of Mount Nimba, scientists from three continents have 
studied the birds. In this way Mount Nimba has become the ornithologically most thoroughly 
explored lowland rain-forest area of Africa. 

The book offers a comprehensive synthesis of information on the avifauna of Mount Nimba 
and its ecological setting. During the 20 years period of biological investigations at Nimba this in 
1962 intact area was gradually opened up by man with far-reaching environmental consequences 
for the rain-forest habitats and profound effects on the birds. Therefore, the book provides not 
only a source of reference material on the systematics, physiology, ecology and biology of the 
birds of Mount Nimba and the African rain-forest, but also data on biogeography in the African 
context as well as on conservation problems. Also behaviour and migration are discussed. At 
Nimba a number of migrants from Europe and/or Asia meet Afrotropical migratory and 
sedentary birds. 

Professor Kai Curry-Lindahl has served as Chairman of the Nimba Research Laboratory and 
Committee since its inception in 1962. Peter Colston is from the Subdepartment of Ornithology, 
British Museum (Natural History), Tring, and Malcolm Coe is from the Animal Ecology 
Research Group, Department of Zoology, Oxford. 

1986, 129pp. Hardback. 565 00982 6 17.50. 






Titles to be published in Volume 52 



Miscellanea 

A revision of the Suctoria (Ciliophora, Kinetofragminophora) 5. The Paracineta 
and Corynophora problem. By Colin R. Curds 

Notes on spiders of the family Salticidae 1. The genera Spartaeus, Mintonia and 
Taraxella. By F. R. Wanless 

Mites of the genus Holoparasitus Oudemans, 1936 (Mesostigmata: Parasitidae) in 
the British Isles. By K. H. Hyatt 

The phylogenetic position of the Yugoslavian cyprinid fish genus Aulopyge Heckel, 
1841, with an appraisal of the genus Barbus Cuvier & Cloquet, 1816 and the 
subfamily Cyprininae. By Gordon J. Howes 

Revision of the genera Acineria, Trimyema, and Trochiliopsis (Protozoa, 
Ciliophora). By H. Augustin, W. Foissner & H. Adam 

The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) with a 
systematic review, a synopsis of Pipistrellus and Eptesicm, and the descriptions of 
a new genus and subgenus. By J. E. Hill & D. L. Harrison 

Notes on some species of the genus Amathia (Bryozoa, Ctenostomata). By P. J. 

Chimonides 



Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester. Dorset 



Bulletin of the 

British Museum (Natural History) 



Mites of the genus Holoparasitus Oudemans 
1936 (Mesostigmata: Parasitidae) in the 
British Isles 

Keith H. Hyatt 



Zoology series Vol52 No 4 30 April 1987 



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ISSN 0007-1498 Zoology series 

Vol52 No. 4 pp 139-1 64 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD Issued 30 April 1 987 



Mites of the genus Holoparasitus Oudemans, 1936 
(Mesostigmata: Parasitidae) in the British Isles 

Keith H. Hyatt 

Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD 



Contents 



Synopsis 

Introduction .... 

Material 

External morphology 
Summary of classification 
Genus Holoparasitus Oudemans 

Key to species 

Descriptions of species . 
Acknowledgements . 
References 




139 
139 
141 
141 
141 
142 
142 
143 
163 
163 



Synopsis 

The mites of the genus Holoparasitus Oudemans, 1936 occurring in the British Isles and the Channel Islands 
are revised. Five species are recorded of which two, H. lawrencei and H. maritimus, are new to science, and H. 
calcaratus (C. L. Koch, 1839) is recorded for the first time. A neotype is designated for H. calcaratus. Habitat 
and distribution data are given and keys to species for males and females are provided. 



Introduction 

The family Parasitidae comprises two subfamilies, the Parasitinae Oudemans, 1901 and the 
Pergamasinae Juvara-Bals, 1972. Hyatt (1980) has revised the British species of Parasitinae. 
Following Evans and Till (1979) the British Pergamasinae comprises the genera Pergamasus 
Berlese, 1904, Amblygamasus Berlese, 1904, Paragamasus Hull, 1918, Holoparasitus Oudemans, 
1936 and Pergamasellus Evans, 1957. Of these, the first three were included by Bhattacharyya 
(1963) in his revision of the British species of Pergamasus sensu lato, whilst the monotypic genus 
Pergamasellus (of which specimens have so far been found only in two localities in southern 
England) is figured by Evans and Till (1979). The remaining genus, Holoparasitus, is revised in the 
present paper. 

The genus Holoparasitus has been mentioned in the British literature scarcely a dozen times and 
only four authors, Halbert (1915), Hull (1918), Turk and Turk (1952) and Turk (1953), discussed 
their determinations. Halbert (1915) recorded calcaratus Koch, pollicipatus Berlese and inornatus 
Berlese from the west coast of Ireland, but he attached reservations to his identifications. Hull 
(1918) recorded the same three taxa from north-east England. Turk and Turk (1952) recorded 
berlesei Oudemans and commented on the confused synonymy of this species, echoing the remarks 
of Oudemans (1936), whilst Turk (1953), in his 'Synonymic Catalogue of British Acari', listed all 
the above names plus four of Berlese's 'varieties' based on specimens in his own collection. The 
remaining authors have given species determinations without comment. Where these specimens 
are available for study they are referred to under the taxa they are considered to represent. 



Bull. Br. Mus. not. Hist. (Zool.) 52(4): 139-164 



Issued 30 April 1987 



139 



140 



K. H. HYATT 




Fig. 1 The National and Irish Grids, showing to the nearest 10-kilometre square the extent of material 
examined during the preparation of the present work. 



Mead-Briggs and Hughes (1966) recorded Holoparasitus pollicipatus from Cambridgeshire and 
Davis (1963) recorded Holoparasitus sp. from Northamptonshire. Unfortunately none of these 
specimens is available for study. 

The most comprehensive revision of the European species of the genus is by Karg (1971) who 
keys and illustrates nine species of which three, calcaratus, inornatus and stramenti Karg, are 
represented in the British fauna. 



MITE GENUS HOLOPARASITUS 1 4 1 

Material 

This revision is based on the examination of over six hundred specimens. The majority 
were removed from Berlese-funnel extractions during the course of my revision of the British 
Parasitinae (Hyatt, 1980). A few were removed from subsequent samples, mainly from areas of 
northern England and south-west Scotland not previously sampled. With the exception, however, 
of the monotypic genus Pergamasellus, Holoparasitus is not found so abundantly as the other 
genera of Pergamasinae, nor is it apparently so widely distributed. 

Few named specimens were already in the Museum collections and of these the majority were in 
the J. H. Murgatroyd and Harry Britten collections from the New Forest area and the north of 
England respectively. 

The map, figure 1, shows on the 10-kilometre squares of the National and Irish Grids the extent 
of the material from which I have examined Holoparasitus in the British Isles and the Channel 
Islands. Localities for Great Britain are given under the current English and Welsh counties and 
Scottish regions or island areas, but if a record is from only a small part of the area the former 
county name is inserted in parentheses. Additionally, large urban areas, e.g. Manchester and 
London, and islands or individual but prominent localities, e.g. Isle of Wight, Island of Mull, or 
Dungeness, are given as such. 



External morphology 

The external morphology of Holoparasitus is essentially typical of the Parasitidae and the terms 
used in the descriptions follow Evans and Till (1979) and Hyatt (1980). As in the majority of the 
Pergamasinae the dorsal setae of the idiosoma are usually of uniform length and thickness, where- 
as in the majority of species of Parasitinae these setae are heterogeneous. Holoparasitus species are 
usually readily separated from the other pergamasines by being conspicuously more spherical in 
body outline, and even under low magnifications the ventral fusion of the holodorsal and opistho- 
gastric shields in the British species of Holoparasitus sensu stricto is easily seen. The exception to 
this last character is in the subgenera Ologamasiphis Holzmann, 1969 and Heteroparasitus Juvara- 
Bals, 1976 (both as yet not recorded from the British Isles) in which the two shields are not fused, 
although the holodorsal shield is continued ventrally, unlike Pergamasus sensu lato which has the 
holodorsal shield entirely dorsal. 



Summary of classification 

Oudemans (1936) proposed Holoparasitus as a new name for Ologamasus Berlese, 1906 
( = Hologamasus [lapsus] Berlese, 1 892) with the type species Gamasus calcaratus C. L. Koch, 1 839. 
The type species of Ologamasus Berlese, 1888 is, by monotypy, Gamasus aberrans Berlese, 1888, a 
member of the family Rhodacaridae (Ryke, 1962, Lee, 1970). 

The first comprehensive review of the genus Holoparasitus was contained in Berlese's 'Mono- 
grafia del genere Gamasus Latr.' published in 1906. In this work the following ten taxa are 
recognised: Gamasus (Ologamasus) calcaratus Koch, 1839 and its two varieties, excisus Berlese, 
1906 and siculus Berlese, 1906; Gamasus (Ologamasus) inornatus Berlese, 1906; Gamasus 
(Hologamasus) pollicipatus Berlese, 1904 and its five varieties appeninorum, cultriger, excipuliger, 
peraltus, andpseudoperforatus, all Berlese, 1906. 

Oudemans (1936) considered that Berlese's pollicipatus and Koch's calcaratus were both 
synonyms of the earlier Acarus lichenis Schrank, 1781. However, Micherdzinski (1969) considered 
that Oudemans had no real evidence for this, bearing in mind that Schrank's figures were so lacking 
in detail. 

Of the recent authors Holzmann (1969) uses the name Ologamasus Berlese, 1892, with which 
she erroneously considers Holoparasitus to be synonymous. She recognises two subgenera, 
Ologamasus s. str. and Ologamasiphis nov., separated by several characters of the deutonymph 



142 K. H. HYATT 

and the female. In the subgenus Ologamasus s. str. she includes calcaratus Koch, 1839, 
inornatus Berlese, 1906, hemisphaericus Vitzthum, 1923, absoloni Willmann, 1940 and intermedius 
Holzmann, 1969, whilst in Ologamasiphis she includes rotulifer Willmann, 1940 and a new species, 
minimus Holzmann, 1969. 

Micherdzinski (1969) follows Oudemans in his preference for Holoparasitus and divides the 
genus into four species-groups: 1 . calcaratus-group and 2. pollicipatus-group, both based on the 
form of the femoral spurs on leg II of the male, 3. Ologamasiphis-group with H. rotulifer and H. 
minimus following Holzmann, and 4. a group to include females which do not belong to previous 
groups, H. excisus (Berlese) and H. hemisphaericus (Vitzthum). 

Karg (1971), at couplet twelve in his key to females, recognises the subgenus Ologamasiphis 
for three species, minimus Holzmann, coronarius Karg (nom. nov. pro rotulifer Holzmann, 1969, 
non Willmann, 1941) and tirolensis Sellnick, 1968, implying that the preceding species belong to 
Holoparasitus s. str. 

Genus HOLOPARASITUS Oudemans 

Hologamasus Berlese, 1904: 235. Non Berlese, 1892: 62. 
Ologamasus Berlese, 1906: 242. Non Berlese, 1888: 194. 
Holoparasitus Oudemans, 1936: 164. 

TYPE SPECIES: Gamasus calcaratus C. L. Koch, 1839. 

Dorsal and ventral shields of the adults well sclerotised; males with holodorsal and opisthogastric 
shields always fused posteriorly; females with holodorsal and opisthogastric shields fused 
posteriorly in Holoparasitus s. str., but free in the subgenus Ologamasiphis Holzmann. 
Deutonymphs with separate podonotal and opisthonotal shields, not strongly sclerotised. In all 
stages setae of dorsal hexagon, i.e./5, z5 and 76, similar to each other and not differing markedly 
from the remaining dorsal stage which are generally short (not exceeding 50 um) and often 
inconspicuous. Tritosternum of male biramous and sometimes modified, base closely associated 
with genital orifice. Tristosternum of deutonymph and female normal, biramous. Junction 
between sternal and metasternal shields of female clique. Genital shield of female broadly 
pentagonal. Opisthogaster with usually not more than 15 pairs of setae. Setaea/ofpalptrochanter 
bifid and with one or more distinct slender processes; setae al v and al 2 of palp femur entire, 
spatulate or setiform. Corniculi strong, entire or notched internally. Legs of deutonymph and 
female without spurs; only leg II of male spurred. Lobes of pulvilli normal, rounded. 

Key to species 
Males 

1 Apophysis on femur II thumb-shaped, about twice as long as axillary process (Fig. 2G) . . 2 
Apophysis on femur II short, hemispherical and not extending beyond the blunt axillary process 

(Fig. 80) 3 

2 Sternogenital shield with a conspicuous 'excipulum' medially between coxae II and HI (Fig. 2B); 

majority of dorsal setae extremely short (c. 10-20 um) (Fig. 2A); idiosoma 590-635 um 

... . . . . . Holoparasitus calcaratus (C. L. Koch) (p. 143) 

Sternogenital region without such a median 'excipular' mark; dorsal setae generally exceeding 
25 um in length; idiosoma 520-570 um . . . . Holoparasitus stramenti Karg (p. 146) 

3 Anterior margin of Sternogenital shield strongly concave medially to behind sternal setae I; a 

conspicuous and shallow transverse structure line present at sternal setae II (Fig. 8B); idiosoma 

530-590 um . . Holoparasitus inornatus ( Berlese) (p. 151) 

Anterior margin of Sternogenital shield not strongly concave medially; sternal ornamentation 
otherwise; idiosoma exceeding 680 um 4 

4 Large species - idiosoma 780-840 um; sternal region with a light but strongly procurved line 

reaching forward from sternal setae II (Fig. 1 OB); tectum trispinate, centre spine long (Fig. IOC); 

corniculi smooth Holoparasitus lawrencei sp. n. (p. 1 55) 

Smaller species - idiosoma 680-750 um; sternal region with reticulations only; tectum broadly 
triangular, granular (Fig. 13C); corniculi cleft on inner margins (Fig. 13G) 

i Holoparasitus maritimussv. n. (p. 158) 



MITE GENUS HOLOPARASITUS 143 

Females 

1 Genital shield produced anteriorly to a tongue-shaped apex (Fig. 6B); endogynium as in figure 6C; 

idiosoma 590-670 urn . . ...... Holoparasitus stramenti Karg (p. 146) 

Genital shield with anterior margin forming an obtuse to right angle, with or without a small 
protruding tip (Fig. 9B) ..... ........ 2 

2 Sternal shield divided longitudinally into two (Fig. 9B); endogynium appears as a circular granular 

structure (Fig. 9C); idiosoma 6 10-690 um . . . Holoparasitus inornatus (Berlese) (p. 151) 
Sternal shield entire (Fig. 11 B) ............. 3 

3 Sternal shield with a pair of indentations on the anterior margin internally to sternal setae I (Fig. 

3B); endogynium as in figure 3C; majority of dorsal setae extremely short (c. 10-25 um) (Fig. 
3A); idiosoma 670-720 jam ..... Holoparasitus calcaratus (C. L. Koch) (p. 143) 
Sternal shield with anterior margin smooth, without indentations ...... 4 

4 Sternal shield granular, with a distinct median pattern, pre-sternal shields entire or divided, 

strongly denticulate (Fig. 14B); endogynium as in figure 14C; idiosoma 770-810 urn 

........... Holoparasitus maritimus sp. n. (p. 158) 

Sternal shield smooth, no distinct median pattern; pre-sternal shields entire or divided, granular 

(Fig. 1 IB); endogynium as in figure 1 1C; idiosoma 850-920 um 

. .......... Holoparasituslatvrenceisp.n.(p.\55) 

Descriptions of species 

Holoparasitus calcaratus (C. L. Koch) 
(Figs 2A-G, 



Gamasus calcaratus C. L. Koch, 1839: Heft 26, Taf. 6. 
Gamasus timidulus C. L. Koch, 1839: Heft 26, Taf. 7. 
Gamasus (Ologamasus) calcaratus: Berlese, 1906: 245. 
Holoparasitus calcaratus: Micherdzinski, 1969: 354. 
Ologamasus (Ologamasus) calcaratus: Holzmann, 1969: 47. 
Holoparasitus excipuliger: Karg, 1971: 361. Non Berlese, 1906. 
Ologamasus pollicipatus Berlese, 1904; 1913: 203 (in part). 

Designation of a neotype 

Oudemans (1906, 1929, 1936) opined that Gamasus timidulus C. L. Koch (?) and Gamasus 
calcaratus C. L. Koch (^) were junior synonyms of Acarus lichenis Schrank, 1781. However, 
Schrank's and Koch's specimens are no longer in existence and although Micherdzinski ( 1 969) and 
Karg (1971) have accepted that timidulus is a synonym of calcaratus, they have not accepted the 
validity of lichenis. Since Oudemans (1936) it has been universally accepted that calcaratus is 
the type of Holoparasitus, but, unfortunately, authors' concepts of calcaratus have not been 
consistent. Specimens labelled Ologamasus pollicipatus in the set of slides accompanying the 
Acarotheca Italica (Berlese, 1913) in the BM(NH) collection are referable to the present species, 
whereas the specimens similarly labelled in the set in the Oudemans collection are rotulifer 
Willmann, 1940. 

Through the courtesy of Dr L. van der Hammen I have been able to examine material from 
several places very close to Regensburg, the type locality. Of the 15 samples of Gamasina 
examined, seven contained Holoparasitus and of these six contained specimens of H . calcaratus. 
One sample contained a single male and female of another species. 

A female from Dechbetten, 1-5 miles (2km) west of Regensburg, Bavaria, 19 July 1959, in 
rotting dry litter, L. van der Hammen coll., is designated as the neotype of Gamasus calcaratus C. L. 
Koch. The specimen is deposited in the Rijksmuseum van Natuurlijke Historic, Leiden, together 
with deutonymphs, males and other females from the same sample. Three males, three females and 
one deutonymph from this sample are retained in the British Museum (Natural History) through 
the courtesy of Dr van der Hammen. 

MALE. The holodorsal and opisthogastric shields, which are fused ventrally posterior to coxae IV, 
are strongly sclerotised and completely reticulated (Figs 2A, B). The dorsum (Fig. 2A) measures 
590-635 um long x 430-480 um wide and bears over 50 pairs of simple setae ranging in length from 



144 



K. H. HYATT 




Fig. 2 Holoparasitus calcaratus (C. L. Koch), male - A dorsum, B venter, C tectum, D chelicera, E palp 
trochanter, femur and genu, F venter of gnathosoma, G leg II. 

35 urn (the vertical setae, y'l) to 10 um or less for the majority in the opisthonotal region. The setae 
are not arranged entirely symmetrically. 

The tritosternum comprises two slender pilose laciniae that arise from below the anterior margin 
of the genital lamina (Fig. 2B). The sternogenital region is strongly reticulated and bears medially 
between coxae II and III a conspicuous oval mark (the 'excipulum' of Berlese). The sternogenital 



MITE GENUS HOLOPARASITUS 



145 




Fig. 3 Holoparasitus calcaratus (C. L. Koch), female - A dorsum, B venter, C endogynium, D tectum, E 
chelicera, F palp trochanter, femur and genu, G venter of gnathosoma. 

and median opisthogastric setae are up to 40 nm in length, whilst the three anal setae and those 
situated posteriorly in the opisthogastric region are as little as 10 um. The stigma is situated 
opposite the posterior margin of coxa III and the peritreme extends anteriorly to coxa I. 

The tectum is shown in figure 2C. The median portion is broad and blunt in all specimens 
examined and the lateral teeth vary. The chelicera is shown in detail in figure 2D. The movable digit 



146 K.H.HYATT 

is 75 um long and bears one large tooth and 5-6 small, sometimes irregular, teeth. The fixed digit 
bears one sometimes blunt tooth towards the tip and up to seven very small teeth in the region of 
the pilus dentilis. The palp trochanter, femur and genu are shown in figure 2E. The anterolateral 
setae of the femur and genu are spatulate, that on the genu is more slender and is bifurcate near to 
its tip. The corniculi (Fig. 2F) are strong and stalked and all the gnathosomal setae are simple. Leg 
II is shown in detail in figure 2G. The femoral apophysis is thumb-shaped and is about twice as long 
as the axillary process. The ventral process on the genu is short and rounded whilst that on the tibia 
is elongate and bean-shaped. All leg setae are slender whilst a number on the tarsi of legs II-IV are 
stronger and pilose on one margin. The ambulacra are normal. 

FEMALE. The holodorsal and opisthonotal shields, which are strongly sclerotised and completely 
reticulated, are fused ventrally anterior to the anus (Figs 3A, B). The dorsum (Fig. 3A) measures 
670-720 (im long x 500-560 |im wide and bears up to 57 or more pairs of simple setae, about 20 
pairs being in the podonotal region and from 34-37 pairs in the opisthonotal region. The vertical 
setae,y7, are the longest and measure c. 40 urn, whilst the shortest setae, the majority of those in the 
opisthonotal region, measure as little as 10 urn and are extremely fine. 

The tritosternum (Fig. 3B) has a narrow base and the paired laciniae are simple. The presternal 
shields are fused to form a single narrow bar. The ventral shields are strongly reticulated and all 
setae are simple. The sternal shield has a pair of indentations on the anterior margin inwards from 
sternal setae I. The reticulations show a bold procurved transverse line extending from posterior to 
coxae II through sternal pores II. The genital shield measures 1 30-1 60 um long x 210-220 urn 
wide. Its anterior margin is formed into a slight obtuse angle or right angle and bears a slender 
pointed tip. The endogynium is shown in figure 3C and in the specimens examined shows con- 
stancy in form despite clearing for examination. The opisthogastric region bears 8-9 pairs of setae. 
The longest of the ventral setae are sternal setae II and III which measure up to 75 |im. The shortest 
ventral setae are around the posterior margin and measure 10 uin or less. In most of the specimens 
examined the outline of the deutonymphal anal shield is retained. The stigma is situated opposite 
the posterior margin of coxa III and the peritreme extends anteriorly to coxa I. 

The tectum (Fig. 3D) has the centre portion tapered and sinuous. The chelicera is shown in figure 
3E. The movable digit measures 87 urn and bears three prominent teeth and two small teeth, whilst 
the fixed digit bears five teeth decreasing in size towards its tip. The palp trochanter, femur and 
genu are shown in figure 3F. The anterolateral seta of the genu is bifurcate whilst those of the tibia 
are spatulate. The venter of the gnathosoma is shown in figure 3G. The internal posterior hypo- 
stomatic setae are the longest (70 um or more) and with the palpcoxal setae are pilose, at least on 
one margin. The hypognathal groove shows only about seven rows of denticles. The leg setae are 
slender and a few on the tarsi are pilose on one margin. The ambulacra are normal. 

MATERIAL EXAMINED. 14 samples -7^^, 169$. 

ENGLAND: Hampshire, Surrey, Sussex/Kent border near the coast, Cambridgeshire (including 
Huntingdonshire). 
This species is recorded mainly from grasses and leaf-litter in damp habitats. 

As far as I can establish, there are no previous authenticated records from the British Isles. 

Holoparasitus stramenti Karg 
(Figs 4A-F, 5A-G, 6A-G) 

Holoparasitus stramenti Karg, 1971: 356. 

Gamasus (Ologamasus) calcaratus var. excisus Berlese, sensu Halbert, 1915: 54 (in part). 

Gamasus (Ologamasus) pollicipatus Berlese, sensu Halbert, 1915: 55. 

Holoparasitus pollicipatus (Berlese) sensu Browning, 1956: 386. 

DEUTONYMPH. The lightly reticulated dorsal shields are weakly sclerotised and yellowish-brown in 
colour. The podonotal shield averages 235 um long x 325 um wide. It bears 17 pairs of simple 
slender setae, none longer than 30 um (Fig. 4A). The opisthonotal shield averages 180um 
long x 225 um wide and bears 12 pairs of simple slender setae from 20-25 um in length. The 
surrounding membrane bears dorsally on each side about 20 short, simple setae. 



MITE GENUS HOLOPARASITUS 



147 




Fig. 4 Holoparasitus stramenti Karg, deutonymph - A dorsum, B venter, C tectum, D chelicera, E palp 
trochanter, femur and genu, F venter of gnathosoma. 



The tritosternum has a narrow base and pilose laciniae. The sternal shield (Fig. 4B) averages 
1 80 um long and is lightly reticulated and weakly sclerotised. The sternal setae average 25 urn long 
and are simple. Pre-sternal shields absent. The oval anal shield bears the usual three setae, each 
simple and about 12 ^im in length. The stigma is situated opposite the anterior margin of coxa IV, 
and the granular peritreme and peritrematal shield extend anteriorly to the level of coxa I. The 1 1 to 
15 or more pairs of simple opisthogastric setae are slender. 

The triangular tectum is serrated anteriorly and bears a slender tip (Fig. 4C). The chelicerae are 
as in figure 4D, the movable digit measures 58 um in the figured specimen. The palp trochanter, 
femur and genu are shown in figure 4E. The anterolateral setae of the femur and genu are spatulate. 
The corniculi and the venter of the gnathosoma are as in figure 4F, the gnathosomal setae being 
simple with the internal hypostomatics the longest. The hypognathal denticles are not discernible. 
All leg setae are slender, the majority are simple but some, especially on tarsus II, are finely pilose 
on one margin. The pulvilli are normal, rounded and with two claws. 



148 



K. H. HYATT 



r tT^ ^ . 



7. v \i. -VM 




Fig. 5 Holoparasitus stramenti Karg, male -A dorsum, B venter, C tectum, D chelicera, E palp 
trochanter, femur and genu, F venter of gnathosoma, G leg II. 

MALE. The holodorsal and opisthogastric shields, which are strongly sclerotised and completely 
reticulated, are fused ventrally anterior to the anus (Figs 5A, B). The dorsum (Fig. 5A) measures 
520-570 um long x 330-400 um wide, is finely granular and bears 45 pairs of simple setae, 20 pairs 
in the podonotal region and 25 pairs in the opisthonotal region. The vertical setae, jl, are the 
longest, measuring up to 45 um. The remaining setae are 20-25 um long. 

The tritosternum comprises two slender laciniae which arise from below the anterior margin of 
the genital lamina (Fig. 5B). The sternogenital region is reticulated and the pattern is constant for 



MITE GENUS HOLOPARASITUS 1 49 

the species. The sternogenital setae and the median opisthogastric setae are the longest - up to 
45 um - whilst the remainder are considerably shorter. The three anal setae are simple. The stigma 
is situated opposite the posterior margin of coxa IV and the peritreme extends to coxa I. 

The tectum (Fig. 5C) is normally symmetrically formed and is trispinate. The chelicera is shown 
in figure 5D; the movable digit is 50 um long in the figured specimen and bears a single tooth near 
the tip, whilst the fixed digit bears two prominent teeth and about four smaller ones. The palp 
trochanter, femur and genu are shown in figure 5E. The anterolateral setae of the femur and genu 
are spatulate. The venter of the gnathosoma is as in figure 5F. The corniculi are stalked and strong. 
The gnathosomal setae are simple and the hypognathal groove bears about nine rows of denticles. 
Leg II is shown in detail in figure 5G. The femoral apophysis is strong and thumb-like whilst the 
axillary process is short, but elongate. The ventral process on the genu is swollen, whilst that on the 
tibia is not prominent. All the leg setae are slender, some on the tarsi being pilose on one margin. 
The ambulacra are normal, with rounded pulvilli and two claws. 

FEMALE. The holodorsal and opisthogastric shields, which are strongly sclerotised and completely 
reticulated, are fused ventrally anterior to the anus (Figs 6A, B). The dorsum (590-670 um 
long x 410-475 um wide) is finely granular and bears 48-49 pairs of simple setae (Fig. 6A), 20 pairs 
in the podonotal region and 28-29 pairs in the opisthonotal region. The vertical setae, jl, are the 
longest, measuring up to 40 um. The remaining setae are about 20-25 um in length. 

The tritosternum (Fig. 6B) has a narrow base and pilose laciniae. The presternal shields are 
fused to form a strong transverse bar, occasionally with a small section at each end being almost 
or entirely detached. The ventral shields are completely reticulated and are granular. The 
reticulations on the sternal shield follow a definite, but simple, pattern with a pair of oblique lines 
originating from the angle between coxae II and III and passing through sternal pores II, but 
petering out before reaching the centre of the shield. The genital shield measures 130-183 um 
long x 190-225 um wide, its size appearing to be related to the actual size of the specimen. The 
genital shield is unique among the British species of the genus in that it is produced anteriorly into a 
strong tongue-shaped apex. The endogynium is shown in figure 6C; its general content appears 
constant, but the position of the 'teeth' varies considerably due to distortion in life or during 
preparation for examination. The opisthogastric region bears 8-9 pairs of setae. The sternal setae 
are the longest - c. 45 um - and the metasternal, genital and opisthogastric setae decrease slightly 
in length in that order. The three anal setae are short (c. 12 um) and simple. The stigma is situated 
opposite the posterior margin of coxa III and the peritreme extends anteriorly to the level of coxa I. 

The tectum (Fig. 6D) is similar to that of the male, but in some specimens irregularity is present. 
The chelicera is as in figure 6E. The movable digit measures 82 um in the figured specimen and 
bears three teeth, whilst the fixed digit bears up to six less prominent teeth. The palp trochanter, 
femur and genu are shown in figure 6F. The anterolateral setae of the femur and genu are spatulate. 
The venter of the gnathosoma is shown in figure 6G. The palpcoxal setae are pilose on one margin 
whilst the other gnathosomal setae are simple. The hypognathal groove bears about eight rows of 
denticles. Some of the leg setae are finely pilose on one margin. The ambulacra are normal. 

MATERIAL EXAMINED. 93 samples - 3 DNN, 80 & 178 ??. 

ENGLAND: Isles of Scilly, Devon, Somerset, Berkshire, Hampshire, Surrey, Sussex, Kent, 
Middlesex, Hertfordshire, Buckinghamshire, Gloucestershire, Bedfordshire, Norfolk, Suffolk, 
Cambridgeshire (including Huntingdonshire), Hereford and Worcester, Leicestershire, 
Cheshire, North Yorkshire, Cumbria (Cumberland, Westmorland), Northumberland. 
SCOTLAND: Lothian/Borders, Strathclyde (Argyllshire), Tayside (Perthshire), Inner Hebrides 
(Mull, lona, Eigg). 

WALES: Gwynedd (Caernarvonshire), Dyfed (Cardiganshire), Clwyd, Gwent, Glamorgan. 
IRELAND: Galway, Mayo, Clare, Leitrim. 
CHANNEL ISLANDS: Jersey. 

Although this species has been collected from leaf-litter, compost, moss and grassland, it has a 
marked preference for wet, marshy habitats. One female from Co. Mayo, Ireland, was taken on the 
seashore. 



150 



K. H. HYATT 




Fig. 6 Holoparasitus stramenti Karg, female -A dorsum, B venter, C endogynium, D tectum, 
E chelicera, F palp trochanter, femur and genu, G venter of gnathosoma. 



DISTRIBUTION. Karg (1971) stated that the holotype of this species came from the Baltic coast and 
gave its distribution as central Europe. The name has not appeared subsequently in the literature. 
H. stramenti is one of the two most abundant British members of the genus. It was recorded from 
Co. Mayo, Ireland, by Halbert (1915) as Gamasus (Ologamasus) calcaratus var. excisus Berlese 
from Achill Island and Westport, and as Gamasus (Ologamasus) pollicipatus Berlese from Clare 
Island. Two females of this species were recorded from Jersey by Browning (1956) as Holoparasitus 
pollicipatus (Berlese). A single undetermined female was recorded from Moor House National 
Nature Reserve, Westmorland, by Block (1965), whilst Davis (1970) recorded a single female from 
Monks Wood National Nature Reserve as Holoparasitus ?pollicipatus. 



MITE GENUS HOLOPARASITUS 1 5 1 

Holoparasitus inornatus (Berlese) 
(Figs 7A-F, 8A-G, 9A-G) 

Gamasus (Ologamasus) inornatus Berlese, 1906: 257. 

Holoparasitus inornatus: Schweizer, 1961: 34 ($ only), Micherdzinski, 1969: 366. 

Ologamasus (Ologamasus) inornatus'. Holzmann, 1969: 47. 

Holoparasitus calcaratus (Koch, 1839) sensu Schweizer, 1961: 36 (J only). Karg, 1971: 361. 

Gamasus (Ologamasus) calcaratus Koch, 1839 sensu Halbert, 1915: 54 (part). 

DEUTONYMPH. The dorsal shields are light yellowish-brown in colour, weakly sclerotised and 
faintly reticulated (Fig. 7A). The podonotal shield averages 460 um long x 300 um wide when 




Fig. 7 Holoparasitus inornatus (Berlese), deutonymph - A dorsum, B venter, C tectum, D chelicera, 
E palp trochanter, femur and genu, F venter of gnathosoma. 



1 52 K. H. HYATT 

flattened and depending on one or more of the marginal (r-series) setae being on or off the shield, 
bears 16-17 pairs of fine simple setae, none measuring more than about 35 um. The opisthonotal 
shield averages 180 um long x 210 um wide and bears 12 pairs of simple setae up to c. 25 um in 
length. All the dorsal setae taper extremely finely. The surrounding membrane bears dorsally on 
each side up to about 20 short simple setae. 

The tritosternum has a narrow base and pilose laciniae. The sternal shield (Fig. 7B) measures 
160-170 um long and is lightly reticulated and weakly sclerotised. The setae are simple. Presternal 
shields absent. The oval anal shield bears the usual three setae, each simple and about 12-15 um in 
length. The stigma is situated opposite the anterior margin of coxa IV and the granular peritreme 
and peritrematal shield extend anteriorly to the level of coxa I. The opisthogastric setae, which 
number about 1 7 pairs, are fine and simple. 

The tectum (Fig. 7C) is triangular and is flanked by small tooth-like projections each side. The 
chelicera is shown in figure 7D; the movable digit measures 72 um in the figured specimen and bears 
three teeth, whilst the fixed digit bears about five teeth. The palp trochanter, femur and genu are 
shown in figure 7E. The anterolateral seta of the femur is broad and pectinate on one margin and 
the two anterolateral setae of the genu are spatulate. The corniculi and venter of the gnathosoma 
are shown in figure 7F. The four pairs of gnathosomal setae are simple whilst the hypognathal 
denticles are indistinct. All leg setae are slender, the majority are simple, but a few on tarsus II are 
finely pilose on one margin. The ambulacra are normal, with rounded pulvilli and two claws. 

MALE. The holodorsal and opisthogastric shields are fused ventrally posterior to coxae IV and are 
heavily sclerotised (Figs 8A, B). The dorsum, which measures 530-600 um long x 390-460 um 
wide, is strongly granular with almost no trace of reticulations (Fig. 8A). It bears, apparently, a 
fairly constant and symmetrically arranged number of setae, 20 pairs in the podonotal region and 
30-31 pairs in the opisthonotal region. The longest setae, the verticals (/7), measure c. 30 um and 
the tendency is for the setae to decrease in length towards the posterior of the dorsum where some 
are as short as 10 um. 

The tristosternum comprises a short base and two pilose laciniae (Fig. 8B). The sternogenital 
region is strongly reticulated and characteristically shaped. The anterior margin of the sterno- 
genital shield is recessed deeply to accommodate the genital lamina and immediately posterior to 
sternal setae II there is a strong procurved ridge right across the shield. These two formations give 
this anterior region a very characteristic appearance. The posterior two-thirds of the sternogenital 
region are characteristically ornamented and are clearly separated from the opisthogastric region 
level with the posterior third of coxae IV, only the endopodal shields retaining their fusion. The 
sternogenital and opisthogastric setae are up to 50 um in length, whilst the three anal setae and 
those around the posterior margin of the ventral surface are about 18 um long. The stigma is 
situated opposite the posterior margin of coxa III and the peritreme extends anteriorly to coxa I. 

The tectum is shown in figure 8C. The broad central part is acutely tapered and is flanked on each 
side by a short prong. The chelicera is shown in figure 8D. The movable digit is 83 um long and 
bears four or five small teeth. The fixed digit bears six to eight small teeth. The palp trochanter, 
femur and genu are shown in figure 8E. The anterolateral setae of the genu and femur are spatulate. 
The corniculi (Fig. 8F) are strong and stalked and are cleft to a varying degree on their inner 
margins. The palpcoxal setae are finely plumose, the remaining three pairs of gnathosomal setae 
are simple. There are about 10 rows of hypognathal denticles. Leg II is shown in detail in figure 8G. 
The apophysis on femur II is short, hemispherical, and does not extend beyond the tip of the 
similarly shaped axillary process. The ventral processes on the genu and tibia are smooth and 
elongate. All leg setae are slender, some on tarsi II-IV are finely pilose on one margin. The 
ambulacra are normal. 

FEMALE. The holodorsal and opisthonotal shields are fused ventrally posterior to coxae IV and 
are heavily sclerotised (Figs 9 A, B). The dorsum (Fig. 9A), which measures 61 0-690 um 
long x 420-570 urn wide, is strongly granular and, like the male, has almost no trace of reticulation. 
It bears about 50 pairs of simple setae, up to 23 pairs in the podonotal region and up to 27 pairs in 
the opisthonotal region. Setae//, the verticals, are the longest, measuring about 35 um and the 
remainder are slightly shorter with a minimum length of c. 18 um posteriorly. 



MITE GENUS HOLOPARASITUS 



153 




Fig. 8 Holoparasitus inornatus (Berlese), male - A dorsum, B venter, C tectum, D chelicera, E palp 
trochanter, femur and genu, F venter of gnathosoma, G leg II. 



The tritosternum has a narrow base and pilose laciniae (Fig. 9B). The presternal shields are 
fused into a single narrow bar. The ventral shields are strongly reticulated and all setae are simple. 
The sternal shield is finely granular and is divided longitudinally at the centre. Additionally, a 
conspicuous procurved line passes through sternal pores II and spans the entire shield. The 
genital shield measures 135-160 urn long x 210-260 um wide. Its apex is formed almost as a right 



154 



K. H. HYATT 




Fig. 9 Holoparasitus inornatus (Berlese), female -A dorsum, B venter, C endogynium, D tectum, 
E chelicera, F palp trochanter, femur and genu, G venter of gnathosoma. 

angle and it does not bear an elongate tip. The endogynium appears to comprise simply an oval 
punctate area with a central detail (Fig. 9C). The opisthogastric region bears 8-9 pairs of simple 
setae. The longest of the ventral setae are the sternals which measure c. 65 um and the shortest are 
around the posterior margin and measure c. 12 um. The stigma is situated opposite the posterior 
margin of coxa III and the finely granular peritreme extends anteriorly to coxa I. 

The tectum (Fig. 9D) has the centre portion finely tapered and it is flanked on each side by a 
single prong. The chelicera is shown in figure 9E. The movable digit measures 95 um in the figured 
specimen and bears three teeth, one large and two smaller but of equal size. The fixed digit bears 



MITE GENUS HOLOPARASITUS 1 5 5 

four teeth, the two distals being small, the two proximals larger. The palp trochanter, femur and 
genu are shown in figure 9F. The anterolateral seta on the femur is pilose distally whilst the two on 
the genu are spatulate. The venter of the gnathosoma is shown in figure 9G. The corniculi are 
strongly formed and the four pairs of gnathosomal setae are simple. Ten rows of hypognathal 
denticles are visible. The leg setae are slender, some on tarsi II-IV are pilose on one margin. The 
ambulacra are normal. 

MATERIAL EXAMINED. 74 samples - 4 DNN, 85 <$<$, 225 ??. 

ENGLAND: Cornwall, Devon, Dorset, Hampshire, Sussex, Kent, Hertfordshire, Gloucestershire, 
Suffolk, Cambridgeshire (including Huntingdonshire), Norfolk, Herefordshire, Warwick- 
shire, Nottinghamshire, Lincolnshire, Cumbria (Westmorland), N. Yorkshire, Durham, 
Northumberland. 

SCOTLAND: Strathclyde (Argyllshire, Mull, Ulva), Tayside (Perthshire), Highland (Inverness- 
shire, Ross and Cromarty), Sutherland, Shetland (Fair Isle). 

WALES: West Glamorgan, Dyfed (Cardiganshire), Gwynedd (Caernarvonshire, Anglesey), 
Clwyd (Denbighshire). 
IRELAND: Leitrim, Mayo, Sligo, Clare, Kerry. 

This species is recorded mainly from mosses, litter and soil in damp habitats and is one of the two 
most abundant representatives of the genus in the British Isles. 

DISTRIBUTION. The only previous records from the British Isles that I have been able to trace and 
authenticate are of specimens recorded by Halbert (191 5) as Gamasus ( Ologamasus ) calcaratus (in 
part) from Co. Mayo and by Davis (1970) as Holoparasitus ?inornatus from Huntingdonshire. 
Davis' (loc. cit.) single female of ?pollicipatus is Holoparasitus stramenti Karg. 

It is recorded from France (Berlese, 1916), Germany (Berlese, 1 906, Karg, 1971) and Switzerland 
(Schweizer, 1961). 

X . 1^ 

Holoparasitus lawrencei sp. nov. 
(FigslOA-G, 11A-G) 

MALE. The holodorsal and opisthogastric shields, which are fused ventrally posterior to coxae IV, 
are strongly sclerotised and completely reticulated (Figs 10A, B). The dorsum (Fig. 10A) measures 
780-840 urn long x 550-610 um wide and bears around 50 pairs of simple setae that range in length 
from 63 um (/7) to 18um in the opisthonotal region. The figured specimen measures 820 um 
long x 610 um wide and bears apparently 46 setae on the left side of the dorsum and 51 on the right 
side. As can be seen from the figure, the setae are not entirely arranged symmetrically. 

The tritosternum comprises two slender pilose laciniae that arise from below the anterior margin 
of the genital lamina (Fig. 10B). The sternogenital region is strongly reticulated and bears a 
strongly procurved line between coxae II and a similar, but less conspicuous, line between coxae III 
and IV. The sternogenital and median opisthogastric setae are up to 65 um in length, whilst the 
three simple anal setae and the posterior ventral setae are about 20 um long. The stigma is situated 
opposite the posterior margin of coxa III and the strongly granular peritreme extends to coxa I. 

The tectum (Fig. IOC) is trispinate and the centre prong is long and sinuous and may be broken 
off. The chelicera is shown in figure 10D. The movable digit is 92 um long in the figured specimen 
and bears no distinct teeth. The fixed digit bears apparently only one rudimentary tooth adjacent to 
the pilus dentilis. The palp trochanter, femur and genu are shown in figure 10E. The anterolateral 
setae of the femur and genu are spatulate, that of the genu is pilose on one margin. The venter of the 
gnathosoma is shown in figure 10F. The corniculi are strong and stalked, the palpcoxal setae are 
pilose whilst the remaining three pairs are simple, and the hypognathal groove bears about eleven 
rows of denticles. Leg II is shown in detail in figure 10G. The femoral apophysis is short and does 
not extend beyond the blunt axillary process. The ventral processes on the genu and tibia are 
shallow and directed anteriorly. All leg setae are slender, a number on the tarsi and tibiae of legs 
II-IV are stronger and pilose on one margin. The ambulacra are normal. 

FEMALE. The holodorsal and opisthonotal shields, which are strongly scelerotised and completely 
reticulated, are fused ventrally anterior to the anus (Figs 1 1 A, B). The dorsum (Fig. 1 1 A) measures 



156 



K. H. HYATT 




Fig. 10 Holoparasitus lawrencei sp. nov., male- A dorsum, B venter, C tectum, D chelicera, E palp 
trochanter, femur and genu, F venter of gnathosoma, G leg II. 

850-920 urn long x 650-710 um wide and bears up to 49 or more pairs of simple setae, about 20 
pairs in the podonotal region and from 29-31 pairs in the opisthonotal region. The figured 
specimen -the holotype - measures 870 urn long x 660 urn wide and bears 35 setae on the 
podonotum, 1 7 on the left and 1 8 on the right, and 60 setae on the opisthonotum, 3 1 on the left and 
29 on the right. The vertical setae,y7, and setaey'2 are the longest, measuring 50 urn or more, whilst 



MITE GENUS HOLOPARASITUS 



157 




Fig. 11 Holoparasitus lawrencei sp. nov., female -A dorsum, B venter, C endogynium, D tectum, 
E chelicera, F palp trochanter, femur and genu, G venter of gnathosoma. 

the shortest setae, the majority of those in the opisthonotal region, measure as little as 12 \am and 
are extremely fine. 

The tritosternum (Fig. 1 IB) has a narrow base and pilose laciniae. The presternal shields are 
coarsely granular and may be entire or divided medially. The ventral shields are reticulated and all 
setae are simple. The sternal shield bears a pair of lines originating from the angle between coxae II 
and III and passing through sternal pores II and almost meeting at the centre of the shield. The 
genital shield measures 170-180 urn long x 265-280 um wide. In the figured specimen- the holo- 
type - it measures 170 um x 280 um. Its anterior margin is formed into slightly more than a right 



1 58 K. H. HYATT 

angle and does not have an extended tip. The endogynium is shown in figure 1 1C. It appears to 
distort easily. The opisthogastric region bears 8-10 pairs of setae. The longest of the ventral setae 
are probably sternal setae II, being up to 75 urn, whilst the shortest, in the opisthogastric region, 
measure only c. 12 um. The three simple anal setae are also short. In the type the post-anal seta is 
bifid. The stigma is situated opposite the posterior margin of coxa III and the peritreme extends to 
the level of coxa I. 

The tectum (Fig. 1 ID) is very similar to that of the male. The chelicera is as in figure 1 IE. The 
movable digit measures 125 um in the figured specimen and bears three blunt teeth, whilst the fixed 
digit bears five blunt teeth. The palp trochanter, femur and genu are shown in figure 11F. The 
anterolateral seta of the femur is broad with one edge pectinate, whilst those of the genu are 
spatulate. The venter of the gnathosoma is shown in figure 1 1G. The setae are simple, and the 
hypognathal groove bears ten rows of denticles. The majority of the leg setae are fine and simple, 
but some of the distal setae on tarsi II-IV are pilose on one margin. The ambulacra are normal. 

MATERIAL EXAMINED. 16 samples - 20 <3<$, 16 $?. 

ENGLAND: Cornwall - Hayle, the holotype $ (1984.12.4. 1) collected by Mr P. N. Lawrence from 
dry, light, leaf-litter, 24.5.1975: Lelant, St Ives, 1 3 in a carrion trap on salt marsh and 1 $ in a 
garden trap, 1943 (Dr F. A. Turk) (these specimens not included in the type series); Isles of 
Scilly-St Agnes, 1 $ paratype (1984.12.4. 9) from litter under Pittosporum, 5.11.1959 (K. H. 
Hyatt); Somerset - Bath, Kennet and Avon Canal, \<S paratype (1984.12.4. 16) from moss, 
humus, etc., 10.3.1962 (P. N. Lawrence); Hampshire - Milford-on-Sea, 1 9 paratype (1984.12.4: 
30) with no data (A. S. Hirst): Isle of Wight, 1 paratype (1984.12.4. 3) with no habitat data, 
April 1948 (T. A. Lloyd); Oxfordshire - Oxford, 1 cJ, 1 $ paratypes (1984.12.4. 17-18) from the 
nest of blackbird Turdus merula, August 1979 (Miss A. Warburton); Norfolk - Blackborough, 1 
cJ paratype (1984.12.4. 2) with no habitat data, 25.2.1969 (Miss A. Reeve); Suffolk - Westleton 
Heath, 3 33 paratypes (1984. 12.4. 13-1 5) from algae on rotten wood, 7.3. 1964 (P. N. and Mrs K. 
Lawrence); Lincolnshire - no locality, 1 ? paratype (1925.6.24. 584) with no habitat data, 1900 
(C. F. George); Cumbria (Lancashire) - Grange-over-Sands, 3 33, 2 $? paratypes (1984.12.4. 
4-8) from tree-holes, 27.1.1954 (D. Macfarlane); Cumbria (Cumberland) - Newton Arlosh, 
Carlisle, 1 3, 2 $$ paratypes (1973.28) with no data (J. E. Hull). 

SCOTLAND: Tayside (Perthshire) - Glen Farg, 2 33 paratypes ( 1 984. 12.4.1 9-20) from mosses on 
deciduous trees, 24.9.1982 (K. H. Hyatt); Dumfries and Galloway (Wigtownshire) - Moss of 
Cree, 5 33, 4 ?? paratypes (1984.12.4. 21-29) from moss in birch tree-holes, 16.9.1982 (K. H. 
Hyatt). 

WALES: Dyfed (Cardiganshire) - Dol-y-Bont, 1 ? paratype (1984.12.4. 12) from damp moss, 
1 5.8. 1 957 (Dr G.O.Evans). 

IRELAND: Clare- Lough Inchiquin, 1 (J, 1 $ paratypes (1984.12.4. 10-1 1) from litter near a weir, 
June/July 1971 (P. N. Lawrence). 

This species is named after Mr P. N. Lawrence whose diligent collecting of soil arthropods has 
done much to increase our knowledge of the British and Irish faunas. 

Holoparasitus maritimus sp. nov. 
(Figs 12A-F, 13A-H, 14A-G) 

Holoparasitus calcaratus: Browning, 1956: 386, Non Koch, 1839. 

DEUTONYMPH. The dorsal shields are pale yellowish brown in colour, lightly sclerotised and 
reticulated (Fig. 12 A). The podonotal shield measures 260-280 um long x up to 480 um wide, 
depending on the degree of lateral displacement of the posterior region as shown in the figure. The 
figured specimen bears essentially 18 pairs of setae although the lefty'2 is missing. Setae jl, the 
verticals, measure c. 35 um whilst the remainder reduce in length to c. 18 um on the margins. The 
opisthonotal shield measures 1 90-240 um long x 260-290 um wide and bears 13 pairs of setae, 
although in the figured specimen the right Z3 is missing. The setae range in length from 1 8 22 um. 
The surrounding membrane bears dorsally on each side from 30-40 fine setae not exceeding 1 8 um 
in length. 



MITE GENUS HOLOPARASITUS 



159 







Fig. 12 Holoparasitus maritimus sp. nov., deutonymph - A dorsum, B venter, C tectum, D chelicera, 
E palp trochanter, femur and genu, F venter of gnathosoma. 

The tritosternum has a narrow base and simple laciniae. The sternal shield (Fig. 12B) measures 
160-170 |im long and is lightly sclerotised and entirely reticulated. The setae are simple. Presternal 
shields absent. The oval anal shield bears the usual three setae and is reticulated. The stigma is 
situated opposite the anterior margin of coxa IV and the granular peritreme and irregularly 
outlined peritrematal shield extend to coxa I. The opisthogastric setae number upwards of 16 pairs 
depending on the actual position of those towards the posterior margin. 



160 K. H.HYATT 

The tectum (Fig. 12C) is essentially triangular and bears strong lateral teeth, but is irregularly 
outlined. The chelicera is shown in figure 12D. The movable digit measures 78 um in the figured 
specimen and bears three widely spaced teeth. The fixed digit bears five or six smaller teeth. The 
palp trochanter, femur and genu are shown in figure 12E. The anterolateral setae of the femur and 
genu are spatulate. The corniculi and the venter of the gnathosoma are shown in figure 12F. The 
anterior and the internal posterior hypostomatic setae are simple whilst the external hypostomatic 
and the palpcoxal setae are lightly pilose on one margin. About seven rows of hypognathal 
denticles are present. All the leg setae are slender and the majority are simple, but a few on tarsus II 
are finely pilose on one margin. The ambulacra are normal, with rounded pulvilli and two claws. 

MALE. The holodorsal and opisthogastric shields are fused ventrally posterior to coxae IV. They 
are heavily sclerotised and entirely reticulated (Figs 13 A, B). The dorsum (Fig. 13 A) measures 
680-750 jim long x 425-500 um wide and bears about 60 pairs of simple setae that range in length 
from c. 55 um (setaey'7) to 1 2 um in the opisthonotal region. The figured specimen measures 730 um 
long x 450 um wide and bears apparently 60 pairs of setae on the left side and 58 pairs on the right 
side. The podonotal region bears 19 pairs of setae arranged symmetrically, whilst the opisthonotal 
region bears 41 setae on the left side and 39 on the right side. 

The tritosternum comprises two slender pilose laciniae that arise from below the anterior margin 
of the genital lamina (Fig. 13B). The anterior margin of the sternogenital shield is moderately 
recessed medially. The ornamentation of the sternogenital region is without a characteristic 
pattern. The sternogenital setae are about 60 um in length whilst the opisthogastric setae are 
shorter. The three anal setae are simple and like those in the posterior region of the opisthogastric 
shield measure approximately 18 um. The stigma is situated opposite the posterior margin of coxa 
III and the peritreme extends anteriorly to coxa I. 

The tectum is strongly granular and produced normally into a triangular process (Fig. 13C). 
However, in the figured specimen it is irregularly formed as shown in figure 13D. The chelicera is 
shown in figure 13E. The movable digit is 97 um long and bears one large tooth and four to five 
small teeth. The fixed digit bears up to seven very small teeth. The palp trochanter, femur and genu 
are shown in figure 1 3F. The anterolateral seta of the femur is broad and slightly pectinate on one 
margin and the two anterolateral setae of the genu are spatulate. The corniculi (Fig. 13G) are 
stalked and are deeply cleft on their inner margins. The gnathosomal setae are all simple and there 
are up to 13 rows of hypognathal denticles. Leg II is shown in figure 13H. The apophysis on femur 
II is short, hemispherical, and does not extend beyond the tip of the swollen axillary process. The 
ventral processes on the genu and tibia are smooth and elongate. All leg setae are slender, some on 
tarsi II-IV are finely pilose on one margin. The ambulacra are normal. 

FEMALE. The holodorsal and opisthonotal shields are strongly sclerotised and fused ventrally 
anterior to the anus (Figs 14A, B). The dorsum (Fig. 14A) is reticulated except in the median 
podonotal region where it is strongly granular. It measures 770-810 um long x 520-560 um wide 
and bears up to about 60 pairs of simple setae, about 20 pairs in the podonotal region and up to 40 
pairs in the opisthonotal region. The figured specimen -the holotype - measures 800 um 
long x 530 um wide and bears 20 pairs of setae in the podonotal region, whilst in the opisthonotal 
region there are 39 setae on the left side and 34 on the right. The vertical setae,y'7, are the longest, 
measuring c. 55 um, whilst the remainder reduce gradually in length towards the posterior of the 
dorsum where the shortest are c. \ 5 um. 

The tritosternum has a narrow base and pilose laciniae (Fig. 14B). The presternal shields are 
fused medially and are strongly denticulate over most of their surface. The sternal shield bears a 
characteristic ornamentation which shows up clearly in alchohol under low magnification. There 
is a longitudinal median design and two pairs of liniae which form part of the reticulation. 
The anterior-most lines run almost diagonally from the centre of the shield towards the 
anterior corners, whilst the second pair runs from the centre through sternal pores II. The genital 
shield measures 160-165 um long x 240-260 um wide. In the figured specimen - the holotype -it 
measures 165 um x 250 um. Its anterior margin is almost right-angled medially and forms a short 
broad tip. The endogynium is shown in figure 14C and appears to distort easily. The opisthogastric 



MITE GENUS HOLOPARASITUS 



161 




Fig. 13 Holoparasitus maritimus sp. nov., male - A dorsum, B venter, C, D tectum, E chelicera, F palp 
trochanter, femur and genu, G venter of gnathosoma, H leg II. . 

region bears 8-9 pairs of setae. The three anal setae are short (c. 18 um) and similar in length to the 
posterior-most ventral setae. The stigma is situated opposite the posterior margin of coxa III and 
the peritreme extends anteriorly to coxa I. 

The tectum (Fig. 14D) is granular and produced into a strong median spine and small lateral 
spines. The chelicera is shown in figure 14E. The movable digit measures c. 108 um long and bears 



162 



K. H. HYATT 




r > t 
/ f V 1 


\ 


j 




'' .'T 

V ' t > 


f 


\T' 1 


\ 


A', ', 


r 



rt^ 







Fig. 14 Holoparasitus maritimus sp. nov., female -A dorsum, B venter, C endogynium, D tectum, 
E chelicera, F palp trochanter, femur and genu, G venter of gnathosoma. 



three strong teeth. The fixed digit bears at least six smaller teeth. The palp trochanter, femur and 
genu are shown in figure 14F. The anterolateral setae on the femur and genu are spatulate, that on 
the femur being pilose on one margin. The venter of the gnathosoma is shown in figure 14G. The 
external posterior hypostomatic setae are simple, the remaining three pairs are lightly pilose. There 
are up to 1 1 rows of hypognathal denticles. The majority of the leg setae are fine and simple, but 
some on tarsi II-IV are pilose on one margin. The ambulacra are normal. 



MITE GENUS HOLOPARASITUS 1 63 

MATERIAL EXAMINED. 8 samples - 6 DNN, 16 <$<$, 20 $9- 

ENGLAND: Cornwall - Kelsey Head, the holotype ? (1984.12.4. 31) and 4 DNN, 8 &J, 16 ?? 
paratypes (1984.12.4. 32^3) from thrift Armeria maritima, etc., on cliffs, 20.5.1975 (P. N. 
Lawrence coll.): Land's End, 1 ? paratype (1984.12.4. 46) from mossy turf, 26.5.1975 (P. N. 
Lawrence): Phillack, Hayle, 2 ?? paratypes (1984.12.4. 47^8) from dried seaweed, 24.5. 1975 (P. 
N. Lawrence): Porthleven, 2 DNN, 3 ^ paratypes (1984.12.4. 49-53) from mossy cliff turf, 
29.5.1975 (P. N. Lawrence); Isles of Scilly-St Agnes, 2 $$ paratypes (1984.12.4. 44-45) from 
thrift and grasses on rocks by seashore, 7.4.1957 (K. H. Hyatt). 

SCOTLAND: Inner Hebrides - lona, 1 $ paratype (1984.12.4. 55) from sandy beach grass with 
Fucus, 3.6.1970 (P. N. Lawrence). 

CHANNEL ISLANDS: Jersey - Elizabeth Castle, 1 $ paratype (1954.3.19. 49) from vegetation on 
cliff-face, 30.8.1951 (Dr G. O. Evans). This specimen was recorded by Browning (1956) as 
Holoparasitus calcaratus (C. L. Koch). 



Acknowledgements 

Of the many collectors who have donated specimens to the British Museum (Natural History) my special 
thanks go to Mr P. N. Lawrence, formerly in the Department of Entomology. 

Colleagues in other institutions have kindly loaned material: Dr L. van der Hammen, Rijksmuseum van 
Natuurlijke Historic, Leiden (Oudemans), Dr M. V. Hounsome, Manchester Museum (Britten) and Dr J. P. 
O'Connor, National Museum of Ireland, Dublin (Halbert). Dr F. A. Turk, Camborne, Cornwall, kindly 
loaned specimens from his own collection. Miss A. S. Baker compared my drawings with material in the 
Berlese collection in Florence and Mr K. P. Martyn prepared the distribution map. 



References 

Berlese, A. 1888. Acari Austro-Americani quos collegit Oloysius Balzan. Manipulus primus. Boll. Soc. ent. 
i/o/. 20: 17 1-222. 

1892. Acari, Myriopoda et Scorpiones hucusque in Italia reperta. Or do Mesostigmata (Gamasidae). 
Padova, 143 pp. 

1904. Acari nuovi. Manipulus I. Redia 1: 235-252. 

1906. Monografia del genere Gamasus Latr. Redia 3: 66-304. 

1913. Acarotheca Italica. Firenze, Fasc. II: 202-204. 
1916. Centuria seconda di Acari nuovi. Redia 12: 125-177. 



Bhattacharyya, S. K. 1963. A revision of the British mites of the genus Pergamasus Berlese s. lat. Bull. Br. Mus. 

nat. Hist. (Zool.) 2: 131-242. 
Block, W. C. 1965. Distribution of soil mites (Acarina) on the Moor House National Nature Reserve, 

Westmorland, with notes on their numerical abundance. Pedobiologia 5: 244-251. 
Browning, E. 1956. On a collection of Arachnida and Myriapoda from Jersey, Channel Islands, with a check 

list of the Araneae. Bull. a. Soc.jersiaise 16: 377-394. 
Davis, B. N. K. 1963. A study of micro-anthropod communities in mineral soils near Corby, Northants. J. 

Anim.Ecol.32:49-ll. 

1970. Some Acarina from Monks Wood National Nature Reserve. Entomologist's mon. Mag. 105: 
220-223. 

Evans, G. 0. 1957. An introduction to the British Mesostigmata (Acarina) with keys to families and genera. J. 
Linn. Soc. Zool. 43: 203-259. 

& Till, W. M. 1979. Mesostigmatic mites of Britain and Ireland (Chelicerata: Acari - Parasitiformes). 
Trans, zool. Soc. Land. 35: 139-270. 

Halbert, J. N. 1915. Clare Island Survey, Part 39ii Acarinida: Section II - Terrestrial and marine Acarina. 

Proc. R. Ir. Acad. 31: 45-136. 
Holzmann, C. 1969. Die Familie Parasitidae Oudemans 1901 (Eine systematische Studie aus dem Jahre 1955). 

Acarologie 13: 3-24, 25-55. 

Hull, J. E. 1918. Terrestrial Acari of the Tyne Province. Trans, nat. Hist. Soc. Northumb. 5, 1: 13-88. 
Hyatt, K. H. 1980. Mites of the subfamily Parasitinae (Mesostigmata: Parasitidae) in the British Isles. Bull. Br. 

Mus. nat. Hist. (Zool.) 38: 237-378. 



1 64 K. H. HYATT 

Juvara-Bals, I. 1972. Mixogamasus, un nouveau genre de Parasitidae (Acariens: Anactinotriches) de 
Romanic. Acarologia 14: 3-14. 

1976. Sur le genre Holoparasitus Oudemans et sur certains caracteres morphologiques de la famille 
Parasitidae Oudem. (Parasitiformes). Acarologia 17: 384-409. 

Karg, W. 1971. Acari (Acarina), Milben Unterordnung Anactinochaeta (Parasitiformes). Die freilebenden 
Gamasina (Gamasides), Raubmilben. Tierwelt Dtl. 59, 475 pp. 

Koch, C. L. 1839. Deutschlands Crustaceen, Myriapoden und Arachniden. Regensburg, Heft 26, no. 6. 

Lee, D. C. 1970. The Rhodacaridae (Acari: Mesostigmata); classification, external morphology and distri- 
bution of genera. Rec. S. Aust. Mus. 16 (3): 1-219. 

Mead-Briggs, A. R. & Hughes, A. M. 1966. Records of mites and lice from wild rabbits collected throughout 
Great Britain. Ann. Mag. not. Hist. (13), 8: 695-708. 

Micherdzinski, W. 1969. Die Familie Parasitidae Oudemans 1901 (Acarina, Mesostigmata). Krakow 
(Pahstwowe Wydawnictwo Naukowe), 690 pp. [German. Polish and Russian summaries]. 

Oudemans, A. C. 1901. Notes on Acari, Ser. 3. Tijdschr. ned. dierk. Vereen. 7: 50-88. 

- 1929. Kritisch Historisch Overzicht der Acarologie. II, 1759-1804. Tijdschr. Ent. 72, Suppl.: 1-1097. 

1936. Kritisch Historisch Overzicht der Acarologie (Critico-historical survey of Acarology). Leiden 
(Brill), IIIA, 1805-1 850, 430 pp. 

Ryke, P. A. J. 1962. The subfamily Rhodacarinae with notes on a new subfamily Ologamasinae (Acarina: 

Rhodacaridae). Ent. Ber., Amst. 22: 155-162. 

Schrank, F. von P. von. 1781. Enumeratio Insectorum Austriae Indigenorum. Augustae Vindelicorum, 548 pp. 
Schweizer, J. 1961. Die Landmilben der Schweiz (Mittelland, Jura und Alpen). Parasitiformes Reuter. 

Denkschr. schweiz. naturf. Ges. 84: 1-207. 
Sellnick, M. 1968. Zwei neue Pergamasus-Arten aus Osterreich. Ber. naturw.-med. Ver. Innsbruck 56: 

463^72. 
Turk, F. A. 1953. A synonymic catalogue of British Acari. Ann. Mag. nat. Hist. (12) 6: 1-26, 81-99. 

& Turk, S. M. 1952. Studies on Acari. 7th series: Records and descriptions of mites new to the British 
fauna, together with short notes on the biology of sundry species. Ann. Mag. nat. Hist. (12) 5: 475-506. 

Vitzthum, H. v. 1923. Acarologische Beobachtungen. 7. Reihe. Arch. Naturgesch. 89A, 2: 97-181. 
\\illmann, C. 1940. Neue Milben aus Hohlen der Balkanhalbinsel, gesammelt von Prof. Dr K. Absolon, 
Brunn (2. Mitteilung). Zoo/. Anz. 130: 209-218. 

1941. Die Acari de Hohlen der Balkanhalbinsel. (Nach dem Material der "Biospeologica balcanica".) 
Studie Oboru vseol. kras. Nauky B. Biol. Ser. 8 (14), 80 pp. 




Peter R. Colston & Kai Curry-Lindahl 



For evolution and speciation of animals Mount Nimba in Liberia, Guinea and the Ivory Coast is 
a key area in Africa representing for biologists what the Abu Simbel site in Egypt signified for 
archaeologists. No less than about 200 species of animals are endemic to Mount Nimba. Yet, this 
mountain massif, entirely located within the rain-forest biome, is rapidly being destroyed by 
human exploitation. 

This book is the first major work on the birds of Mount Nimba and surrounding lowland 
rain-forests. During 20 years (1962-1982) of research at the Nimba Research Laboratory in 
Grassfield (Liberia), located at the foot of Mount Nimba, scientists from three continents have 
studied the birds. In this way Mount Nimba has become the ornithologically most thoroughly 
explored lowland rain-forest area of Africa. 

The book offers a comprehensive synthesis of information on the avifauna of Mount Nimba 
and its ecological setting. During the 20 years period of biological investigations at Nimba this in 
1962 intact area was gradually opened up by man with far-reaching environmental consequences 
for the rain-forest habitats and profound effects on the birds. Therefore, the book provides not 
only a source of reference material on the systematics, physiology, ecology and biology of the 
birds of Mount Nimba and the African rain-forest, but also data on biogeography in the African 
context as well as on conservation problems. Also behaviour and migration are discussed. At 
Nimba a number of migrants from Europe and/or Asia meet Afrotropical migratory and 
sedentary birds. 

Professor Kai Curry-Lindahl has served as Chairman of the Nimba Research Laboratory and 
Committee since its inception in 1962. Peter Colston is from the Subdepartment of Ornithology, 
British Museum (Natural History), Tring, and Malcolm Coe is from the Animal Ecology 
Research Group, Department of Zoology, Oxford. 

1986, 129pp. Hardback. 565 00982 6 17.50. 






Titles to be published in Volume 52 

Miscellanea 

A revision of the Suctoria (Ciliophora, Kinetofragminophora) 5. The Paracineta 
and Corynophora problem. By Colin R. Curds 

Notes on spiders of the family Salticidae 1. The genera Spartaeus, Mintonia and 
Taraxella. By F. R. Wanless 

Mites of the genus Holopamsitus Oudemans, 1936 (Mesostigmata: Parasitidae) in 
the British Isles. By K. H. Hyatt 

The phylogenetic position of the Yugoslavian cyprinid fish genus Aulopyge Heckel, 
1841, with an appraisal of the genus Barbus Cuvier & Cloquet, 1816 and the 
subfamily Cyprininae. By Gordon J. Howes 

Revision of the genera Acineria, Trimyema, and Trochiliopsis (Protozoa, 
Ciliophora). By H. Augustin, W. Foissner & H. Adam 

The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) with a 
systematic review, a synopsis of Pipistrellus and Eptesicus, and the descriptions of 
a new genus and subgenus. By J. E. Hill & D. L. Harrison 

Notes on some species of the genus Amathia (Bryozoa, Ctenostomata). By P. J. 

Chimonides 



Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 



Bulletin of the 

British Museum (Natural History) 



The phylogenetic position of the 
Yugoslavian cyprinid fish genus Aulopyge 
Heckel, 1841, with an appraisal of the genus 
Barbus Cuvier & Cloquet, 1816 and the 
subfamily Cyprininae 



Gordon J. Howes 



Zoology series Vol52 No 5 28 May 1987 



The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four 
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and 
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London SW7 5BD, 
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Zool.) 



Trustees of the British Museum (Natural History), 1987 



The Zoology Series is edited in the Museum's Department of Zoology 

Keeper of Zoology : MrJ. F. Peake 
Editor of Bulletin : Dr C. R. Curds 
Assistant Editor : Mr C. G. Ogden 



ISBN 565 05029 X 

ISSN 0007 -1 498 Zoology series 

Vol52 No. 5 pp 165-1 96 
British Museum (Natural History) 
Cromwell Road 
London SW75BD Issued 28 May 1987 




The phylogenetic position of the Yugoslavian cyprinid 
fish genus Aulopyge Heckel, 1841, with an appraisal of 
the genus Barbus Cuvier & Cloquet, 1816 and the 
subfamily Cyprininae 

Gordon J. Howes 

Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD 

Contents 

Introduction 165 

Methods and materials 166 

Nomenclature 167 

Abbreviations used in the figures 168 

Anatomical characters in Aulopyge and their phylogenetic 

significance ^r**~-Lr~"J~" ... 169 

169 
174 
177 

Other osteological features . . . ,/. /.'' . '."'. . \ .VN 182 
Sexual dimorphism and genitalia . . ; . / . . . ,,,.,-, V j 186 

Discussion ! I . 38 {1^2937 .| #. J 186 

Aulopyge relationships and barbin classification! . i "~ . '"."j . .j . H 186 
Immediate relationships of Aulopyge . V\ \- | __,,;. ' 7/y 190 

Acknowledgements V\. ^^*s^ <>,. _.. ,] *:**' t - /J 192 

References ' ; -^:^ 1Q7 

IVC1C1 C11CC& ...... ^^S. *"*^ / " f' *^ fi * Sy^ * ^ 

Appendix 1 . The genus Barbus sensu stricto . ^^^^ : - "AL ^&_**\ l >^&r 193 

Definition and included species .... <i! ^ rr:= irrTF' ^=^'^-- *^ . . 193 
The systematic positions of Barbus brachycephalus Kessler, 1872 and B. mursa 

(Giildenstadt, 1773) 195 

The systematic positions of Barbus andrewi Barnard, 1937 and B. serra Peters, 1864 195 

Appendix 2. Characteristics of the subfamilies Cyprininae and Leuciscinae . . 196 

Introduction 

A ulopyge is a monotypic genus represented by the species A.huegeliiHeckel, 1841 (Fig. 1), endemic 
to rivers and lakes of the Yugoslavian karst regions of Dalmatia. Regrettably, there appears to be 
no published information on the ecology of Aulopyge. Populational data are lacking and the 
species is classified as rare (Lelek, 1980: 122). 

Aulopyge possesses several unique characters (detailed below) which have led to its being placed 
in a separate taxonomic category, the Aulopygini (Bleeker, 1863; Karaman, 1971). In Karaman's 
(1971) view Aulopyge represents a relic of an earlier Eurasian barbine assemblage having a close 
relationship with the schizothoracine cyprinids a group now confined to high-Asia. Lelek (1980: 
122) simply comments '. . . it is difficult to compare it with other taxa'. Arai (1982: 146) concluded 
from his study of karyotypes that Aulopyge, which is polyploid (2n = 100), possesses a 'mosaic of 
barbine and gobionine characters'. 

The recent acquisition by the British Museum (Natural History) of well-preserved specimens of 
Aulopyge huegelii makes possible, for the first time, a detailed anatomical study of the species. The 
information gained from this study has provided a basis not only for a discussion of the phylo- 
genetic status of Aulopyge but also of the classification and relationships of the genus Barbus and 
other cyprinines. 



Bull. Br. A/MS. not. Hist. (Zool.) 52(5): 165-196 Issued 28 May 1987 

165 



166 



G. J. HOWES 





Fig. 1 Aulopyge huegelii, female (above) 127 mm SL and male (below) 106 mm SL. 



In an earlier paper (Howes, 1981: 47-49) the classification of barbelled and non-barbelled 
Cyprinidae was discussed. It was reasoned that one lineage, the barbelled cyprinids, could be 
defined on the synapomorphic presence of paired maxillary barbels each associated with a 
maxillary foramen (or its suggested past presence) and a rostrally extended supraethmoid. 
Following the original division of the European Cyprinidae by Bonaparte (1846), the barbelled 
carps were recognised as the subfamily Cyprininae, while the non-barbelled taxa were ranked as 
the subfamily Leuciscinae. The latter group was defined simply by absence of maxillary barbels 
and associated nerve foramen, no synapomorphy having been discovered that would define it as a 
natural group. 

It can be assumed from the outset therefore that Aulopyge belongs to the Cyprininae, since it 
possesses a pair of maxillary barbels supplied by a branch of the VII facial nerve. From this 
standpoint a more refined hypothesis of the relationship between Aulopyge and other cyprinids 
may be attempted. 



Methods and materials 

The osteology of Aulopyge huegelii was studied from an alcian-alizarin stained and a dry skeletal 
preparation (BMNH) 1903.12.4: 41-5), and from X-Radiographs of specimens 106, 112 and 
127 mm SL (BMNH 1985.8.20: 1-3; Busko Lake, south Bosnia). Genital anatomy was studied in 
dissected specimens of this latter series. 

Comparative osteology of a wide range of cyprinoids was studied both from alizarin stained and 
dry skeletal preparations. A principal data source has been the large collection of X-Radiographs 
of cyprinoid specimens in the BMNH. The following list is ofBarbus specimens used in this study. 

A = alizarin stained preparation; D = dissected specimen; S = dry skeleton. All catalogue 
numbers are BMNH. 

Barbus ablabes 1983.3.30: 7-14(D), B. albanicus 1970.9.24: 265-67(D), B. altianalis eduardianus Uncat., (S), 
B. a. radcliffi 1981.4.9: 42-66(D), B. altus 1898.4.2: 196-205(D), B. amphigramma 1980.7.18: 319-332(D), B. 
andrewi 1900.11.6: 58(D), 1903.4.27: 94^95(S), B. antinorii (type) 1908.10.14: 7, B. arabicus 1976.4.7: 



CYPRINID FISH GENUS A ULOPYGE 1 67 

201-272(D), B. argenteus 1907.6.29: 217(S), B.( = Puntius) arulus 1978.8.31 : 234-259(D), B. aspilus 1909.4.29: 
74(S), B. barbulus 1974.2.22: 1275-77(A), B. barbus 1864.4.11: 41-42(D), 1908.12.28: 123(S), 1985.10.16: 
62-ll(A),\9S5.\Q.l6:5l,B.(^Puntius)bimaculatusl9Sl.\.2l:209-2\l(D),B.binotatus\9S43.3:39-60(D), 
1970.9.3: 56-85(A), B. biscarensis 1970.3.1: 100-125(D,A), B. bocagei 1980.8.20: 1-6(D), B. brachycephalus 
1899.7.25: 25-27, B. burmanicus 1894.5.21: 46-55(D), B. bynni 1861.9.9: 65(S), B. callemis 1951.4.10: 
1-20(A,D), 1 869. 1 .29: 4(S), B. camptacanthus Uncat. (S), B. canis 1 974.2.22: 1292-94(D), Uncat., (S), B. chola 
1935.10.18: 32^6(D), B. ( = Puntius) collingwoodi 1892.9.2: 52-56(A), 1982.4.21: 37-38(D), B. comiza 
(syntype) 1909.7.29: 1, B. conchonius 1978.8.31: 21-35(D), B. cummingi 1978.8.31: 186-222(A), B. 
dorsolineatus 1965.3.15: 406-435(D). B. esocinus 1920.3.3: 80-82(D), B. eutaenia 1965.3.15: 93-122(D), B. 
( = Puntius)filamentosus 1981.1.21: 242-260(D), B.fritschi 1904.1 1.28: 59(S), B.graellsii 1908.2.12: 21^9(D), 
B. grahami 1907.5.4: 52-57(D), B. grypus 1920.3.3: 1-18(D), B. guirali 1902.11.12: 119(S), B. haasianus 
1976.3.18: 892-93(A), B. harterti 1902.7.28: 35(S), B. holotaenia 1984.7.5: 22-27(D), B. holubi 1937.10.4: 
12-14(D), B. hospes 1980.7.18: 434-438, B. hypsolepis 1971.11.26: 28^1(A), B. intermedius intermedius 
1974.1.16: 128-162(A), 166-179(D), 1902.12.13: 338(S), intermedius australis 1893.12.2: 36(S), B.johnstoni 
1975.8.3: 576-80, B. kersteni 1978.8.3: 632-84(D), B. ksibi 1934.10.25: 1-14(D), B. leonensis 1974.9.18: 
77-177(A), B. lineomaculatus 1974.1.16: 396-41 1(D), B. litamba 1974.1.11: 88-93(D), B. lithopides 1889.2.1: 
559-61(D), B. longiceps 1936.4.6: 5-11, 1949.9.16: 90-92, 1864.8.20: 21(S), B. luteus 1874.4.28: 23(S), 
1968.12.13: 201-212(D), B. macrolepis 1972.11.28: 9-1 2(D), B. macrops 1960.6.7: 111-160(D), B. mattozi 
1962.8.22: 2-6(D), B. meridionalis 1935.10.28: 14-17(D,S), B. minimus 1974.1.16: 276-292(D), B. mursa 
1872.5.30: 67-68, B. nasus 1902.1.4: 22(S), B. natalensis 1862.8.28: 8(S), B. neglectus 1980.7.10: 1-26(D), B. 
neumayeri 1969.3.6: 31-50(D), B. ( = Puntius) orphoides 1974.10.10: 865-872(D), B, oxyrhynchus 1893.12.2: 
31(D), 1906.8.25: 17(S), B. paludinosus 1979.3.1: 1-53(D), 1908.1.20: 84(S), cf. paludinosus Uncat., (A), B. 
paytoni 1976.2.2: 29-31(D), B. ( = Puntius) pentazona 1954.11.23 7-82(A), B.perince 1907.12.2: 1268-77(D), 
B. plebejus plebejus 1887.4.5: 15-16, 1982.2.24: 149-1 55(D), plebejus peloponnesius 1964.6.12: 20-26(D), B. 
poechi 1962.7.5: 4-15(D), B.progenys 1903.7.28: 155(S), B.profundus 1970.5.14: 19-30(D), B.(=Tor)putitora 
1884.2.1: 52(S), B. radiatus 1982.4.13: 4597^605(D), B. reinii 1903.10.29: 10(S), B. rocadasi 191 1.6.1: 26(S), 
B. rothschildi 1902.7.28: 22-26(D), B. ( = Puntius) sarana 1933.8.19: 7-14(D), B. schejch 1931.12.21: 4(D), B. 
sclateri (syntypes) 1861.11.20: 9-13, B. serra 1937.10.4: 6-ll(D,S), B. setivemensis 1905.11.28: 59(S), B. 
sharpeyi 1920.3.3: 71-75(D), B. ( = Puntius) sophore 1889.2.1: 777-782(D), B. subquincunciatus 1934.10.29: 
1(D), B. ( = Tor) tambroides 1982.4.21: 39(D), B. tenuis 1975.12.29: 250-265(D), B. thalamakanensis 
1976.3.18: 363-550(D), B. ( = Puntius) titteya 1974.6.11: 8-12(A), B. (=Tor)tor 1893.6.30: 31-38(D), B. 
trimaculatus 1907.4.9: 98(S), B. tropidolepis 1936.6.15: 599-629(A), B. xanthopterus 1973.5.21: 198(D). 

Species without a suffix and those cited in the text but not listed above have been examined by 
X-Radiography only. 

Nomenclature 

Because the concept of cyprinid subfamilies and other higher categories used here differs from that 
of previous authors (see Discussion) I have adopted the following nomenclature. 

Subfamily Cyprininae (cyprinines): a monophyletic assemblage (see text) which includes the 
following subgroups: 

*barbins: a possibly monophyletic group, the members of which possess a foraminate dilatator 
fossa (see text and Table 3 for included taxa). This group embraces, in part, the Barbinae and 
Barbini of previous authors. 

*labeins: a monophyletic group sensu Reid, 1982 and 1985; includes Labeinae, Labeini, 
Labeoinae and Garrini of previous authors. 

*squaliobarbins: a monophyletic group sensu Howes, 1981. 

*schizothoracins: a supposed monophyletic group (see text); the Schizothoracinae and 
Schizothoracini of previous authors. 

*other cyprinines: an unresolved assemblage of taxa not included in any of the above categories 
and lacking a foraminate dilatator fossa (see text and Table 3). 

Subfamily Leuciscinae (leuciscines): a possibly non-monophyletic assemblage including 
Abraminae, Cultrinae etc. of previous authors. 

After this paper had been submitted for refereeing, my attention was drawn to a publication by 
Chen et al. (1984). These authors have proposed an hypothesis of cyprinoid relationships whereby 
they recognise the Cyprinidae as comprising two 'series', the Barbini and Leuciscini. They further 
recognise two monophyletic groups (tribes) within the Barbini, viz. Barbines and Tincanes, of 



168 



G. J. HOWES 



which the Tincanae, Cyprininae, Barbinae and Labeoninae (sic) are the constituent lineages. My 
concept of Cyprininae corresponds to Chen et al. 'Barbini', whilst my subgroups embrace their 
subfamilies. 

The appellations 'small' and 'large' are often given to African Barbus species. As used here, 
'small' refers to those species in which the striae on the exposed part of the scale are radiate, the fish 
usually less than 1 50 mm SL adult size, and the body often marked with spots or lateral stripes; 
'large' refers to those species in which the scale striae are more or less parallel, the fish more than 
1 50 mm SL adult size, and the body lacking any noticeable markings. 



Abbreviations used in 



aa anguloarticular 

abr 1 1 st branched anal fin ray 

afsl-3 anal fin rays (unbranched) 

ah anterohyal 

asn anterior supraneural 

at anal tube 

bb basibranchials 

bh basihyal 

bo basioccipital 

bp basioccipital process 

bsr branchiostegal ray 

cb ceratobranchials 

ccf coracoid-cleithral foramen 

cl cleithrum 

cor coracoid 

csi cavum sinus imparis 

ct connective tissue 

de dentary 

df dilatator fossa 

dfo dilatator foramen 

dfs dorsal fin rays 

dh dorsohyal 

eb epibranchials 

ect ectopterygoid 

enf ectopterygoid facet 

ent entopterygoid 

epf entopterygoid-palatine facet 

ep epural 

epo epioccipital 

fc frontal canal 

fm foramen magnum 

fr frontal 

frl frontal lamina 

hb hypobranchial 

hmf hymandibular fossa 

hs haemal spine 

hyo hyomandibula 

hyp hypurals 

hys hypurapophysis 

ic intercalar 

ih interhyal 

int intestine 

io infraorbitals 

iop interoperculum 

ip infrapharyngobranchials 

lac lachrymal 

let lachrymal canal tube 

le lateral ethmoid 



lef 

lien 

loc 

me 

met 

mp 

nc 

nca 

ns4 

nspu 2 

op 

OS 

ov 

pa 

pc 

pel 

pe 

ph 

phy 

po 

poc 

pro 

ps 

pte 

pts 

ptt 

qf 

ra 

rp 

sb 

sec 

scp 

se 

so 

sop 

sor 

sp 

spr 

srp 

sy 

vh 

vo 

I 
II 

V 

VII 
IX 
X 



the figures 

lateral ethmoid facet 

lateral ethmoid-entopterygoid ligament 

lateral occipital fenestra 

mesethmoid 

metapterygoid 

masticatory plate of basioccipital 

neural complex 

neural canal 

neural spine of 4th centrum 

neural spines of 2nd preural centrum 

operculum 

orbitosphenoid 

oviduct 

parietal 

parietal canal 

postcleithrum 

preethmoid 

posterohyal 

parhypural 

preoperculum 

preopercular canal (bone enclosed) 

prootic 

parasphenoid 

pterotic 

pterosphenoid 

posttemporal 

quadrate facet 

retroarticular 

proximal radials 

splenial bone 

subcutaneous canal 

scapula 

supraethmoid 

supraoccipital 

suboperculum 

supraorbital 

sphenotic 

sphenotic process 

supraethmoid rostral process 

symplectic 

ventrohyal 

vomer 

olfactory nerve foramen 

optic fenestra 

trigeminal nerve foramen 

facial nerve foramen 

glossopharyngeal nerve foramen 

vagus nerve foramen 



CYPRINID FISH GENUS AULOPYGE 

-srp 
se 



169 



sor 




exo 



bo 



Fig. 2 Aulopyge huegelii, neurocranium in dorsal (left) and ventral (right) views. Scale bar in mm. 



Anatomical characters in Aulopyge and their phylogenetic significance 

The cranium of Aulopyge is shown in Figs 2 and 3. In general appearance it is depressed and 
elongate. The ethmoid region is narrow and shallow, the supraethmoid bearing a sloped, valley- 
like depression and anteriorly having slight lateral expansions and a short rostral extension which 
is medially indented (srp, Fig. 2). The kinethmoid (Fig. 4d) is of the rod-shaped type considered by 
Howes (1978; 1981) as plesiomorphic for cyprinoids. Each lateral ethmoid is extended medially 
along the parasphenoid and contacts its partner, their being no anterior myodome. Laterally, each 
bone extends a narrow, posteriorly pointing wing which ventrally bears a well-developed round 
facet against which the entopterygoid facet articulates (lef, Fig. 2). This is an unusual feature and is 
discussed further below. 

The frontals are narrowed anteriorly and nasal bones are absent; the supraorbital bones are 
small but not excessively reduced. Otherwise, the cranium of Aulopyge exhibits no features which 
may be regarded as anything but plesiomorphic among cyprinoids, viz.: the prootic is elongate 
with a long lateral commissure, the subtemporal fossa is round and deep, there is no posttemporal 
fossa, and the basioccipital has a short posterior process and small, round masticatory plate 
(Figs 2 & 3). 

Likewise the jaws and elements of the suspensorium (Fig. 4), other than the entopterygoid 
(discussed below), show no departure from the 'generalised' cyprinoid morphology (see Howes, 
1978, 1981, 1984). 

The lateral ethmoid and its articulation with the entopterygoid 

The presence in Aulopyge of a facet, ventrally on the lateral ethmoid, apposing an entopterygoid 
facet is a feature which has a restricted distribution amongst the Cyprinidae. Ramaswami (1955) 
drew attention to a mesial entopterygoid facet articulating with the lateral ethmoid in Labeo 



170 



G. J. HOWES 



so P a 



fc 



epo 



se 




exo 



1C 



loc 



fm 



csi 



Fig. 3 Aulopyge huegelii, neurocranium in lateral (above) and posterior (below) views. Scale bar in mm. 



macrostoma and Cyprinus carpio. Howes ( 1 976: 46) noted that such a facet was variously developed 
in cyprinids, supposing it best developed in those species with a long ethmoid region and least in 
those with a short ethmoid. However, further investigation has not endorsed this claim and it 
appears that the presence of an entopterygoid facet is not positively correlated with the length 
of the ethmoid. Its presence seems to require a purely phylogenetic rather than a functional 
explanation. Thus, entopterygoid-lateral ethmoid facets occur only in taxa included in the 
Cyprininae, being absent, but for a single exception (Tinea; see below), in the Leuciscinae, (i.e. all 
non-barbelled cyprinids). The most highly developed form of this articulation occurs in some 
species of Barbus, Cyprinus and in the schizothoracin genus Diptychus (Figs 5 & 6). 

In Cyprinus, the ventral surface of the lateral ethmoid wing is broadly triangular with the ventral 
articular facet situated antero-medially (Fig. 5b); the facet is sloped posteriorly and articulates 
against a round facet on the dorso-anterior border of the entopterygoid, just posterior to that 
bone's articulation with the palatine. 

In Barbus barbus, B. nasus, B. plebejus, B. bocagei, B. meridionalis and B. barbulus the lateral 
ethmoid facet is a large triangular platform (Fig. 5a). In some 'large' Barbus species, e.g. the Asian, 
B. grahami, Barbus ( = Tor) tor and the North African, B. setivemensis the articular, boss-like 
facet is situated at the midpoint of the lateral ethmoid wing (Figs 6c-e). In all these species the 
entopterygoid facet is moderately developed. In yet other African and Asian 'large' Barbus species 
the lateral ethmoid facet lies along the posterior margin of the wing and in some taxa, e.g. the 
majority of 'large' African Barbus and Varicorhinus species, a distinct facet is barely developed, 
there being only a bevelling of the posterior border of the wing (Figs 5d & 6f). In these taxa an 
entopterygoid articulatory surface is feebly developed also (Fig. 5d). However, in the majority of 
African and Asian Barbus examined lateral ethmoid and entopterygoid facets are lacking. This 
appears to be the condition in all the so-called 'small' African Barbus species. 

Amongst schizothoracins a lateral ethmoid facet is variously developed (Figs 6k-m), but in 



CYPRINID FISH GENUS AULOPYGE 

.hyo met enf 



171 




Fig. 4 Aulopyge huegelii. (Above) suspensorium in lateral view; (below), (a) palatine; (b) maxilla; (c) 

premaxilla; (d) kinethmoid. Scale bar in mm. 



enf 








Fig. 5 Articular facets on the ventral surface of the lateral ethmoid wing and antero-dorsal surface of 
the entopterygoid in: (a) Barbus barbus; (b) Cyprlnus carpio; (c) Tor putitora', (d) Barbus oxyrhynchus; 
(e) Tinea tinea; (f) Labeo coubie, entopterygoid facet also shown in lateral view. In (a) dashed outline 
represents attachment area of lateral ethmoid ligament. Anterior to the left. Scale bar = 5 mm. 



172 



G. J. HOWES 











n 




Fig. 6 Lateral ethmoid facets of: (a) Diptychus dybowski; (b) Barbus nasus; (c) B. grahami; (d) B. 
setivemensis; (e) B. lithopides; (f) Varicorhinus tanganicae; (g) Barbus callensis; (h) B. serra; (i) B. 
progeny s; (j) B. canis; (k) Schizothorax grahami; (1) S. taliensis; (m) S. intermedius; (n) S. esocinus and S. 
richardsoni. Semi-diagrammatic; all drawn to same scale; anterior to the left. 



none, apart from Diptychus (Fig. 6a) is there a condition approaching that in the Eurasian Barbus 
species cited above, and an entopterygoid facet is rarely present. 

In the squaliobarbins (Squaliobarbus , Ctenopharyngodon and Mylopharyngodori), a group con- 
sidered as primitive cyprinines (see Howes, 1981, and Fig. 21), the lateral ethmoid articular surface 
is elongate, with a bevelled anterior margin against which abuts the posterior edge of the palatine. 
The entopterygoid articulates only with the posterior rim of the lateral ethmoid wing as in some 
'large' African Barbus described above. 

In labeins, Labeo (sensu Reid, 1 985) has an extensive lateral ethmoid whose ventral surface bears 
a fossa which cups an entopterygoid condyle (Fig. 5f). Garra, on the other hand, has a narrow 
lateral ethmoid wing, which is only connected ligamentously with the entopterygoid. 

Lateral ethmoid and entopterygoid facets are also lacking in Cyprinion, Gibelion and Capoeta; 
whether this condition represents secondary loss or a plesiomorphic state is uncertain in the 
absence of recognised synapomorphies indicating the relationships of these taxa. 

That there is a phylogenetic rather than a functional basis for the various types of articulatory 
surfaces among cyprinines is seemingly supported by the following observations. 

In those taxa where there is a well-developed articulation between the two bones, e.g. Cyprinus 
and some Eurasian Barbus species, the anterior portion of the entopterygoid is almost horizontal 
(Figs 8a & b), and it is also horizontal in those taxa which have only a moderate articulation 
between these bones, e.g. some 'large' African Barbus and Varicornhinus species (Fig. 8c). In 
Aulopyge, where there are well-developed lateral ethmoid and entopterygoid facets, the 
entopterygoid slopes at an angle similar to that in taxa which lack such close articulation, e.g. 
Schizothorax esocinus (Fig. 8d). Thus, whilst the slope of the entopterygoid is correlated with 
cranial width (being horizontal in those taxa with the widest crania) there is apparently no corre- 
lation between slope (both in the horizontal and vertical planes) and the presence or absence of 



CYPRINID FISH GENUS AULOPYGE 



173 




lien 



Fig. 7 Connection between the lateral ethmoid and entopterygoid in, (a) Cyprininae; Barbus barbus, 
and (b) Leuciscinae; Raiamas loati. Scale bars = 3 mm. 



articulatory surfaces. Even if one accepts this as evidence for the apomorphic status of articulatory 
facets, there is nothing to suggest which type of facet morphology is the more derived, be the 
extensive well-developed articulation of the Eurasian Barbus and Cyprinus or the less intimate 
connection of the African Barbus and Varicorhinus species. 

It was noted above that all but one leuciscine taxon lack an articular connection between the 
lateral ethmoid and entopterygoid. Instead, the two bones are ligamentously connected and often 
the entopterygoid extends anterior to the lateral ethmoid (Fig. 7b). Tinea is the exception amongst 
leuciscines, in that the entopterygoid bears a distinct and deep fossa which articulates with a lateral 
ethmoid facet (Fig. 5e). 

That articulatory lateral ethmoid and entopterygoid facets occur only amongst cyprinines 
further supports an internal division of the Cyprinidae, but whether this represents the derived 
condition, and if so, whether it is synapomorphic for those taxa in which the articulation occurs is 
problematic (see remarks above). 

The types of ligamentous connection between the lateral ethmoid and the entopterygoid 
support the subfamilial division of the Cyprinidae (see p. 166 and Appendix 2). The widespread 
ostariophysan condition is for there to be a strong ligament running from the upper medial face 
of the lateral ethmoid wing to the dorsolateral surface of the entopterygoid (Fig. 7a; see also 
Vanderwalle, 1977, Fig. 4 of Barbus barbus). 

In all members of the subfamily Cyprininae investigated, apart from Ctenopharyngodon, there is 
a single, slender ligament connecting the bones; in Ctenopharyngodon a broad ligamentous band 
connects the bones. In the subfamily Leuciscinae a ligament of the type found in the Cyprininae is 
absent and connection between the lateral ethmoid and entopterygoid is via undifferentiated 
connective tissue. Vandewalle (1977) showed in Leuciscus leuciscus a ligament (labelled Li 18) 
running from the edge of the lateral ethmoid to the entopterygoid. I find no such discrete ligament, 
but instead thickened connective tissue running to the lateral edge of the entopterygoid (Fig. 7b). 

The widespread occurrence and constant position of the lateral ethmoid-entopterygoid ligament 
amongst ostariophysans suggests it is plesiomorphic and thus its absence in the Leuciscinae is 
considered a derived loss. It is interesting to note in this group, as compared with the Cyprininae, 
what appears to be an anterior shift of the entopterygoid head, and its somewhat looser connection 
with the palatine, features which may be associated with the absence of a ligamentous connection. 



174 



G. J. HOWES 

.enf 




Fig. 8 Anterior views of the right suspensorium in; (a) Cyprinus carpio; (b) Barbus barbus; (c) 
Varicorhinus tanganicae; (d) Schizothorax esocinus; (e) Aulopyge huegelii. Scale bar for a-d = 5 mm, 
for e = 1 mm. 



Sensory canals and their associated bones 

Aulopyge possesses the pattern of supraorbital canals corresponding to Illick's (1956) group 
IVAA, where a marked gap separates the supraorbital and infraorbital canal systems; the frontal 
and parietal canals are distant and the parietal canals are separated from one another by a midline 
gap. The dorso-cranial canals are bony tubes lying on the surface of their respective bones; the 
frontal canal contains 9-10 pores. The infraorbital series is reduced to bony tubes, the first and last 
infraorbitals being fragmented into several elements (Fig. 9a). 

The most unusual feature of the Aulopyge infraorbital canal is its disassociation from the 
lachrymal (1st infraorbital, lac, Fig. 9a). In an alizarin stained specimen of 52mm, the weakly 
ossified canal lies somewhat ventral to the well-developed elongate 'lachrymal' bone. This situ- 
ation is evident in all the specimens of Aulopyge examined (60-127 mm SL) with the exception of an 
84 mm SL female, where a bony canal tube is attached to the face of the lachrymal (Fig. 9b). Those 
portions of the canal posterior and anterior are epidermal. 

The mandibular-preopercular canal is incomplete. Only a single, small tube lies below and 
separate from the dentary (Fig. 4). There is a short groove along the ventro-lateral border of the 
dentary, but no sign of a canal associated with the anguloarticular. The canal reappears as a series 
of weakly ossified, epidermal tubes along the posterior part of the preoperculum; at the point of 
curvature, the canal runs through the bone (poc, Fig. 4), then continues in three or four epidermal 
tubes, the last terminating close to the dorsal tip of the preoperculum. 

The development of cyprinoid sensory canals was studied by Lekander ( 1 949) who summarised 
the results and theories of previous authors. Lekander showed that the sensory canals can, from the 
earliest ontogenetic stages either be united with their respective bone, later fuse with it, or remain 
separate from it. He drew particular attention to the 'antorbital' ( = lachrymal of most authors) 
noting that in some cypriniforms, the canal remains separate from its lamellar portion. Such is the 
case in the Cobitidae, where there is apparently an antorbital, i.e. a bone lying antero-dorsal to the 
1st infraorbital, while the elongate lachrymal is by-passed ventro-laterally by the subcutaneous 
sensory canal (Lekander, 1949; Parshall, 1983). 

As in the adult Aulopyge, the developing infraorbital canals in some cyprinids often appear 
irregularly spaced and remain unfused to one another; a 'splenial' bone may be present (Lekander, 
1 949: 8 1 ), and the preopercular latero-sensory canal tubes remain separated from one another and 
from the preoperculum (Lekander, 1949: 95; 102; 1 12). 



CYPRINID FISH GENUS AULOPYGE 



175 



lac 






let 



Fig. 9 Infraorbital bones of Aulopyge huegelii; (a) complete series of 52 mm SL specimen; (b) the 
lachrymal of an 84 mm SL specimen with canal attached to the bone, (c) Barbus barbulus, showing 
disassociated lachrymal canal. Scale bars in mm. 



Lekander (1 949: 1 1 3) makes the point that sensory canals in cyprinids develop later than in most 
other teleosts he examined. Whether this is so or not, I observe a temporal difference in the 
development of the infraorbital canals between two species of Barbus. In specimens of Barbus cf. 
paludinosus of 17 mm SL the sensory canal of the 1st infraorbital (lachrymal) is present in the bone 
although it does not become completely enclosed until 24 mm SL. However, in Barbus barbus of 
25 mm SL, the canal is subcutaneous and well-separated from the membranodermic part of the 
lachrymal. These species are respectively, tropical and temperate, and small and large sized. Thus, 
the variation in canal development may reflect the different temperature and hormonally con- 
trolled growth rates. In an adult specimen of Barbus barbulus, the posterior part of the canal lies 
subcutaneously, whereas the anterior part is attached to the lachrymal (Fig. 9c). 



176 







Fig. 10 Lachrymal (1st infraorbital) bones of; (a) Barbus barbus of 66 mm SL; (b) B. barbus, adult; (c) B. 
comiza; (d) B. plebejus; (e) B. capita (also in B. sclateri); (f) B. bocagei (also in B. albanicus); (g) 
B. longiceps; (h) B. grypus (also in B. canis, B. sharpeyi, B. reinii); (i) B. intermedius intermedius; (j) B. 
trimaculatus; (k) B. altus; (1) B. callipterus; (m) Labeo coubie; (n) 5. mursa; (o) 5. semz; (p) 5. andrewi. 
Scale bars = 1 mm. 



If Lekander (1949) is correct in recognising three distinct types of association between the 
laterosensory and membranodermic parts of the canal bones (at least amongst cypriniforms), then 
it may be that these represent arrest at successive ontogenetic stages. In this case, that exhibited by 
Aulopyge and some cobitids where the sensory and membranodermic components are separate 
represented the earliest, whilst that in which they are united, as in Leuciscus, would represent the 
most advanced ontogenetic stage. 

The lachrymal in Aulopyge is virtually oblong in lateral view being somewhat tapered anteriorly. 
In most cyprinids the lachrymal is a deep, triangular or pentagonal bone, as in Cyprinus, Labeo and 
the majority of Barbus species (Figs lOi-m). In some Eurasian Barbus species, however, the 
lachrymal has the same oblong shape as in Aulopyge, and the sensory canal also runs in the ventral 
part of the bone. In this latter respect the Eurasian species also differ from other African and Asian 
Barbus where the canal runs centrally through the lachrymal (Fig. 10k). In Barbus barbus, one of 



C YPRINID FISH GENUS A ULOPYGE 1 77 

the species with an oblong lachrymal in adults, there is a marked ontogenetic change in the bone's 
shape. In a specimen of 66 mm SL, it is almost square with a short, dorsally curved sensory canal 
(Fig. lOa). In adults, the bone is elongated, with a greatly lengthened canal (Fig. lOb), the anterior 
part of the canal having become more deeply forked and an additional pore developing at the 
posterior elongation of the canal. 

The adult lachrymal morphology of Barbus barbus resembles that characteristic of certain other 
Eurasian species (Figs 9c, lOc, d, f & g). A variant of this condition is found in the Middle-eastern 
species B. canis, B. sharpeyi, B. grypus, B, reinii and the Asian Barbus (=Tor) tor, where the 
anterior part of the sensory canal runs close to the anterior border of the bone, and the dorsal 
border is concave (Fig. lOh). 

It is difficult to evaluate the shape of the lachrymal as a phylogenetic character. Skelton (1980) 
pointed out that the South African West Cape species Barbus andrewi and B. serra possess a 
lachrymal of the same elongate form as that of the Eurasian species. However, the lachrymal of 
these two species differs from that in the Eurasian taxa in having the ventral border convex rather 
than straight (Figs lOo & p); see also p. 195. 

My own comparisons permit the following generalisations: 

*in all Leuciscinae the lachrymal has a square or even rounded, never elongate shape, even in 
those species with a relatively long ethmoid region (e.g. Elopichthys bambusa; Fig. 12A in Howes, 
1978). 

*in Barbus there is some degree of 'intermediacy' in shape between such forms as B. trimaculatus 
and B. altus (Figs lOj & k) and the B. canis type (Fig. lOh) exemplified by B. oxyrhynchus and B. 
intermedius (Fig. 10i). 

*there is a distinct (?apomorphic) type characterising a group of Eurasian Barbus species; see 
above. 

Vertebral column, dorsal and anal fins 

The general morphology of the Weberian ossicles and centra ofAulopyge resembles that ofBarbus 
barbus. In both taxa the neural complex is low, with a concave anterior border. Its posterior border 
is irregular and widely separated from the 4th neural spine, which is almost half the height of the 
neural complex and is inclined posteriorly. 

Neural complex. The comparative morphology of the cyprinid neural complex has not been 
subject to any detailed treatment and from the following perfunctory observations appears 
worthy of closer study. The so-called 'neural complex' in cypriniforms is a supraneural having 
synchondral contact with the 3rd and 4th neural arches. There is usually a long gap between the 
supraoccipital and the neural complex and only rarely are they in close contact (see Reid, 1985). 
Within the Cyprinidae, two morphotypes of neural complex are recognisable (briefly described in 
Howes, 1981: 29-30; see also Chen et al, 1984); these can be correlated with the subfamily division 
already recognised as Cyprininae and Leuciscinae (see above and Appendix 2). 

In Cyprininae, the neural complex is most often tall, axe-shaped and lamellate, with a vertical or 
forwardly inclined anterior border and without a grooved dorsal surface. The 4th neural spine is 
rarely as high as the neural complex, most often being half or less than half its height and narrowly 
separated from it. The first free supraneural never articulates directly with the neural complex. 

In Leuciscinae, the neural complex is most often low, oblong or square, vertically or backwardly 
inclined; its dorsal surface contains a groove, and in some taxa, the neural complex is deeply 
forked (Howes, 1981, Fig. 22); the 1st free supraneural articulates with the groove (Howes, 1978: 
19; Fig. 13). The 4th neural spine is most often as tall as the neural complex and may be widely 
separated from it. 

The morphology of the neural complex is variable within the Cyprininae, but from the data at 
hand it is possible to make a broad and tentative classification. Within Barbus, the 'small' species 
examined (B. radiatus, B. paludinosus, B. perince, B. leonensis, B. hulstaerti) and some Asian taxa 
(including B. ( = Puntius] sophore} possess a tall, oblong neural complex, either vertical or sloping 
backward and narrowly separated from the 4th neural spine which is the same height as the neural 
complex (Figs 1 lf-1). 



178 



G. J. HOWES 

nc 



ns4 




Fig. 11 Neural complex (shaded) and position of 4th neural spine in (a) Aulopyge huegelii; (b) Barbus 
barbus; (c) B. plebejus; (d) B. altianalis radcliffi; (e) Cyprinus carpio; (f) Barbus paludinosus', (g) 
B. perince; (h) Puntius sophore; (i) Barbus marequensis ('long-head morph'); (j) Varicorhinus 
steindachneri; (k) V. ensifer; (1) Schizopygopsis stoliczkae. Drawings made from radiographs, all to 
approximately the same scale. 



CYPRINID FISH GENUS A ULOPYGE 1 79 

Within the 'large' Barbus species, as in other Cyprininae, the neural complex is tall and axe- 
shaped. Its relationship to the 4th neural spine is variable. In some taxa the spine is short and 
curved forward, e.g. B. intermedius , B. arabicus, B. altianalis (Fig. lid), B. (=Tor) putitora, 
Carassius auratus, or long and curved forward, e.g. Cyprinus carpio (Fig. 1 le), Cydocheilichthys, 
or short, vertical or sloping backward, e.g. majority of 'large' African Barbus (Fig. Hi), and some 
Asian Barbus. In some Varicorhinus species the spine is minute and barely developed as is also the 
case in Cyprinion species (see Howes, 1982). In all these taxa, however, the 4th neural spine is 
closely apposed to the posterior border of the neural complex (Fig. 1 Ij). 

As noted above, the neural complex of Aulopyge huegelli and Barbus bar bus exhibit another 
morphotype (Figs 1 la & b), being squat to oblong with a concave anterior border and an indented 
posterior border leaving a wide gap between it and the 4th neural spine. Other taxa with this 
morphology are the Eurasian, Middle-eastern and Chinese Barbus species plebejus, nasus, 
meridionalis , barbulus, schejch, subquincunciatus and grahami. An exaggerated variant of this 
condition occurs among the schizothoracin genera Schizocypris (Fig. 1 11), Diptychus, Gymnocypris 
and Schizothorax, where the neural complex is irregularly shaped and widely separated from a 
small 4th neural spine. 

From this limited survey it cannot be said which of these is a derived morphotype. That 
characteristic of Aulopyge, some Eurasian and Middle-eastern Barbus and schizothoracins may 
simply be a correlate of the generally elongate and depressed bodies of those taxa. There is also a 
degree of intraspecific and ontogenetic variability. For example, the ontogenetic sequence of 
neural complex development in the 'large' African Barbus intermedius is at 21 -5 mm SL (Fig. 12a) 
that of the adult morphology (almost identical to that of B. altianalis, shown in Fig. 1 Id) in which 
the complex is narrowly separated from the 4th neural spine. At 25 mm SL the neural complex is 
tilted forward, is relatively taller and has a large gap separating it from the neural spine. At 3 1 mm 
SL the neural complex is upright and the 4th neural spine is tall and narrowly separated from it. 

The four ontogenetic stages shown in Fig. 12 of specimens 21-5, 23-5, 25-0 and 31 -0 mm SL seem 
to reflect four of the similar adult morphotypes described above. 

To summarise the conditions of the neural complex among cyprinines: 

*tall and oblong with long 4th neural spine in 'small' African Barbus and (?all) Asian Puntius 

*tall and axe-shaped with 4th neural spine closely apposed in 'large' African and Asian Barbus 
and most other cyprinines, subgrouped as: 

4th neural spine short some African Barbus and other cyprinines 
4th neural spine long most African and Asian Barbus 
4th neural spine minute Varicorhinus and Cyprinion 

*low, oblong or square with irregular anterior and posterior borders and with 4th neural spine 
widely separated posteriorly in Aulopyge, Eurasian Barbus and schizothoracins 

Vertebral number. Aulopyge has a total of 37-38 vertebrae, of which- 10 (including the four 
Weberian vertebrae) are pre-dorsal, i.e. the neural spine of the last vertebra in the set lies in front 
of the 1st dorsal pterygiophore. This total vertebral number lies within the modal range for 
Cyprininae. 

In a sample of 46 'large' African Barbus species the range is 36-42, of which 20 species have a 
range of 9-11 pre-dorsal vertebrae, 4 species have 11-12 (oxyrhynchus , somereni, mariae and 
ethiopicus) and the remaining 22 species have 13-17. These latter species, apart from the South 
African Cape B. serra and B. andrewi, are European and Middle-eastern species (Table 1). 
Schizothoracin genera have both higher total (46-48) and pre-dorsal (13-17) vertebral numbers 
(Table 2). In other Cyprininae, the numbers of pre-dorsal vertebrae rarely exceed 10; in Cyprinion 
there are 8-12, in Cyprinus 9-10, Gibelion 8 and Catlacarpio 8-9. In labeins, Garra has 9-12, and 
Labeo has 8-9. Squaliobarbin taxa also have a high pre-dorsal number, 10-12. 

Skelton (1976) recorded the vertebral numbers in four groups of African Barbus, groupings 
made on the basis of scale striae pattern and degree of ossification in the last unbranched dorsal fin 
ray. He found higher counts in the group with parallel striated scales and with the dorsal fin ray 
ossified and smooth, a group to which belong the 'large' African Barbus species cited above. 



180 



G. J. HOWES 






Fig. 12 Barbus intermedius Ontogenetic development of the neural complex, at (a) 21-5 mm SL; (b) 
23-5 mm; (c) 25 mm; (d) 31 mm. Scale bars = 0-5 mm. 



Following Lindsey's (1976) broader discussion of pleomerism, Skelton (1980) pointed out that 
Jordan's rule (the correlation of increased vertebral number with higher latitudes) may be a factor 
when considering, for example, the endemic high-latitude, high-altitude redfin 'Barbus' which 
have a more frequently occurring range (36-38) than species of 'small' African Barbus (31-38). 
Skelton argues that such specialisation signifies that the higher vertebral number represents a 
synapomorphy, one he uses to recognise the redfin 'Barbus' as a monophyletic group. 

In the Cyprininae, the total vertebral number never exceeds 48, and the modal range is 38-40; in 
the Leuciscinae the total range is greater, being 33-61, as is the modal range of 40-45 (see Howes, 
1978, Table 1). Perhaps more significant is the consistently higher range of pre-dorsal vertebrae in 
Leuciscinae, 10-19 versus 9-14 in Cyprininae. Howes (1978; 1984) considered a high number of 
vertebrae as a synapomorphy for the aspinin group of leuciscine cyprinids, since the range for this 
group exceeds that of other leuciscines in both abdominal and caudal vertebrae. 



CYPRINID FISH GENUS AULOPYGE 



181 



Table 1 Vertebral and lateral line counts in Barbus species having high total and pre-'dorsal numbers of 
vertebrae and lateral line scales, and having a serrated last unbranched dorsal fin ray. In ' Barbus' species with 
a pre-dorsal vertebral count of 9-1 1, the total count rarely exceeds 43. 



Species 


Total 


Pre-dorsal 


Lateral line scales 


albanicus 


44 


14 


57 


andrewi 


38^0 


14-16 


38-40 


barbulus 


44 


13 


52-54 


barbus 


46 


14 


55-63 


bocagei 


42^4 


14-15 


45-49 


brachycephalus 


47 


11 


63 


capita 


42-45 


13 


57-65 


(including specimens labelled as kersiri) 








comiza 


43 


12 


48-50 


(syntypes) 








esocinus 


48 


14 


76-78 


graellsi 


42^3 


14 


47-52 


grypus 


44-47 


13-14 


40 


lacerta 


43 


13 


55-63 


longiceps 


43^44 


13-14 


51-60 


meridionalis 


40 


13 


48-60 


mursa 


43 


14 


90-97 


nasus 


43^4 


13-14 


49-78 


plebejus 


41^2 


13-14 


49-78 


rajanorum 


45-46 


13-14 


57-65 


(including specimens labelled as schejch) 








sclateri 


42 


12 


46-47 


(syntypes) 








serra 


39^1 


14-17 


42^3 


sharpeyi 


40-42 


13-14 


30-31 


(lacks serrated last dorsal spine) 








subquincunciatus 


45 


13-14 


80-84 


xanthopterus 


44 


13 


58-60 



Table 2 Vertebral counts in a selection of schizothoracins. 



Species 



Total 



Pre-dorsal 



Diptychus dybowski 
Diptychus maculatus 
Gymnocypris sp. 
Schizothorax dipogon 
Schizothorax esocinus 
Schizothorax chrysochlorus 
Schizothorax grahami 
Schizothorax intermedius 
Schizothorax richardsoni 
Schizothorax prenanti 
Schizothorax sinuatus 
Schizothorax yunnanensis 
Schizopygopsis stoliczkae 



48 

49 

46 

49 

46-47 

42 

47 

48 

46 

46 

48 

46 

48 



182 



G. J. HOWES 



dfs 
2-3 




Fig. 13 



rp 



Unbranched dorsal fin rays of, (a) Aulopyge huegelii, 1 1 5 mm SL; (b) Barbus barbus, 24 mm SL 
(cartilage stippled); scale = 1 mm. 



It is difficult to assign polarity to vertebral number for other groups of cyprinids because, 

*there is a continuum from the relatively low numbers in Cyprininae to the higher numbers in 
Leuciscinae 

*there is the phenomenon of pleomerism (see Lindsey, 1975) 

*vertebral numbers may be influenced by latitudinal position and temperature changes (see 
Lindsey, 1975; Lindsey &Arnason, 1981). 

Lindsey (1975) commented that the Catostomidae display significant pleomerism among its 
species, but not in the family as a whole. The same observation can be applied to the Cyprinidae, 
where deep-bodied genera such as Cyprinion and Megalobrama have similar maximum lengths to 
those of cylindrical, depressed or compressed and slender forms, but possess lower vertebral 
numbers. 

Dor sal fin and serrated unbranched dorsal fin ray. In Aulopyge the first (reduced) dorsal fin ray lies 
on a vertical just anterior to the base of the pelvic fin and above the 15th vertebra. 

In those Barbus with a high number of pre-dorsal vertebrae the 1st dorsal fin ray lies above the 
16th-18th vertebra and above or somewhat anterior to the origins of the respective fins. In the 
majority of the Cyprininae, the dorsal fin lies: 

*above or anterior to the origin of the pelvic fins. 

*rarely posterior to the pelvic fin origin, (e.g. 'Labeo 1 stoliczkae, Barbus paludinosus, B. serra, 
some Puntius species). 

In the Leuciscinae, however, only exceptionally does the dorsal fin originate in advance of the 
pelvics, (e.g. Pogonichthys). In both Cyprininae and Leuciscinae there are several taxa where the 
dorsal fin origin is immediately above the pelvic fin insertion. Such a situation occurs in both basal 
leuciscines with short, cylindrical bodies, (e.g. Opsariichthys, Zacco) and those with elongate, 
compressed bodies, (e.g. Salmostoma, Macrochirichthys). In cyprinines, this generalised position 
of the dorsal fin is present in many 'large' and 'small' African Barbus species. Skelton (1980) 
considered a dorsal fin posteriorly placed in relation to the pelvics as a synapomorphy uniting 
serrated-dorsal fin rayed redfin Barbus species. However, if one assumed the Cyprininae to be the 
derived sister-group of the Leuciscinae, such a posterior dorsal fin position may indicate the 



CYPRINID FISH GENUS AULOPYGE 



183 



ah 



hb1 



ph 



ih 



bb1-3 




p2-3 



cb4-5 



Fig. 14 Aulopyge huegelii, branchial arches of left side in dorsal view. Scale = 2 mm. 



plesiomorphic condition. The forward placement of the dorsal fin in the Cyprininae, seen in its 
most extreme form amongst labeins, is more likely to be the derived state. 

An analysis of the position of the 1st dorsal fin ray in relation to the vertebral column again 
reflects the major taxonomic grouping of the Cyprininae and Leuciscinae. In the majority of 
cyprinines, the 1st dorsal ray lies above the ll-18th vertebra (14 15th in Aulopyge), whereas in 
leuciscines, it may lie above any from the 16th to the 31st vertebra (modally between the 18th and 
2 1 st). The furthest posterior position of the 1 st dorsal fin ray occurs in the chelin assemblage, where 
it lies above the 21st-26th vertebra in Salmostoma and the 30th-31st in Macrochirichthys. In the 
schizothoracins the 1st dorsal ray lies above the 17th-21st vertebra. 

In Aulopyge there are 3 unbranched dorsal fin rays, the last being moderately serrated along its 
distal posterior border; there are 7-8 branched dorsal fin rays. 

The number of unbranched dorsal fin rays preceding the 1st branched ray varies in the 
Cyprinidae from 2-6. Gosline (1978) found some significance in the numbers of unbranched dorsal 



184 



G. J. HOWES 



cl 



scp 




Fig. 15 Aulopyge huegelii, right pectoral girdle in lateral view. The (medial) positions of the 
mesocoracoid and part of the scapula are indicated by dashed lines. 



fin rays, believing a modal count of 4 to be representative of the Cyprininae whilst 3 was present in 
'. . . other cyprinid subfamilies'. Although Gosline's subfamily concept differs from that presented 
here, I find his statement justified. A possible reason for there being a high number of unbranched 
dorsal fin rays in cyprinines may be correlated with the often marked ossification of the last such 
element. A large heavy spine-like ray, in order to remain rigid may require some anterior bracing in 
the form of several and strong elements in the fin. 

Highly ossified dorsal rays rarely occur in the Leuciscinae, (e.g. Capoetobramd) and never bear 
serrations. 

In the Cyprininae the last unbranched ray is always the largest but varies from flexible to heavily 
ossified, and may be smooth or serrated along its posterior margin. When present, a serrated ray 
may bear serrae over its entire or partial proximal length. A serrated dorsal ray occurs only in some 
species of the genera Barbus, 'Puntius\ Schizothorax and Mystacoleucus, while in other genera, 
such as Acrossocheilus , Cyclocheilichthys and Cyprinus, all species possess a serrated last 
unbranched dorsal ray. 

Based principally on the classification of Boulenger (1911), Skelton (1976) recognised four 
group of Barbus, of which only one (Group III) contained species with a serrated dorsal fin ray. 
Within this group, the subgroup (IIIA) comprises the 'large' African Barbus and contains those 
species which also have relatively high total and pre-dorsal vertebral counts (see above, p. 1 79 and 
Table 1). The ranking of serrated dorsal fin rays as a synapomorphy is dubious since the feature 
has an irregular distribution among genera recognised as monophyletic, (e.g. Cyprinion; see 
Howes, 1982). However, it would be possible to test for the plesiomorphic nature of dorsal fin ray 
serrations by observing their presence in some ontogenetic stage of those taxa whose adults lack 
them. In Barbus barbus, in which the last unbranched dorsal ray bears serrations, they begin to 
appear at 23-5 mm SL when that ray is still segmented (Fig. 1 3b). 



CYPRINID FISH GENUS AULOPYGE 



185 



hyp 6 




Fig. 16 Aulpyge huegelii, (above) caudal fin skeleton of 52 mm SL specimen. Scale = 1 mm; (below) 
variation on second neural spine (dark shading) on PU 2 of specimens (a) 106mm, (b) 112mm, (c) 
127mmSL. 



Other osteological features 

Aulopyge is conservative in its other skeletal elements. 

The gill arches are of a generalised cyprinid type except for the complete absence of gill-rakers on 
the outer margin of the 1st ceratobranchial and only 3 or 4 rakers on the 1st epibranchial. The 
pharyngeal bone (5th ceratobranchial, Fig. 14) is broad and bears a single row of four teeth, the 
first somewhat globular with a prominent cusp, the others having bevelled or chisel-like crowns. 

The pectoral girdle has a tall, upright cleithral limb and a short horizontal limb with a narrow 
lamina (Fig. 1 5). The cleithral-coracoid foramen is minute and the coracoid is small. The size of the 
cleithral-coracoid foramen is variable amongst cyprinids, both intra- and interspecifically (see 
Howes, 1979: 180), and appears to have little worth as a phyletic character. There is a single, long 
postcleithrum in Aulopyge. 

The caudal fin skeleton is of a generalised type with 6 hypurals, a well-developed hypurapophysis, 
paired uroneurals and a long, proximally expanded epural (Fig. 16). There is, however, variable 
development in the neural arch on PU 2 . In the smallest specimen available (52 mm SL) there are 
two neural arches on PU 2 , the posterior arch having only a short spine (Fig. 16). In a specimen of 
106 mm SL there are two arches with fully developed spines, and in the largest, 127 mm SL, the 
condition resembles that of the smallest specimen, namely, the second, posterior arch having a 
small neural spine (Figs 16a-c). 



186 




Fig. 17 Aulopyge huegelii, anal tube and anal fin of female. Drawn from dissection and X-Radiograph 

of 1 16 mm SL specimen. 

Radiographs of a wide range of cyprinines reveal the presence of a second neural arch on PU 2 to 
be of not infrequent occurrence, although when it does occur, the neural spine is usually fully 
developed, (e.g. Barbus plebejus , Barbuscanis, Barbusmicropogon, Barbus barbulus , Acrossocheilus 
yunnanensis, Carassius auratus). A reduced second PU 2 neural spine is found in Barbus comiza. 

The significance, phyletic or otherwise, of a second neural arch and spine on PU 2 is unknown. Its 
mosaic and wide distribution in cyprinines make polarity assignment impossible. It is of interest to 
note, however, that in leuciscines, it is the 3rd preural centrum which bears a double neural arch 
rather than the 2nd as in cyprinines (see Howes, 1984: 296). Variability of neural arches on the 
posterior caudal centra may be a plesiomorphic feature of teleosts; Greenwood (1970: 134) noted 
such variability in Elopiformes. 

Sexual dimorphism and genitalia 

Seeley (1 886) pointed out the marked sexual dimorphism of Aulopyge exhibited in the morphology 
and position of the anal and genital openings and in the smaller body size of the male. 

In the male Aulopyge, the anus and genital opening are separated, the genital orifice being 
posterior in position and lying in front of the first unbranched anal fin ray. In the female, both 
openings and their respective ducts are contained in a fleshy tube which is adnate to the 2nd 
unbranched anal fin ray. The oviduct is firmly joined to the flexible 2nd ray for part of its length 
(Fig. 1 7). In both males and females the 1 st unbranched anal fin ray is vestigial and does not project 
from the body surface. The genital morphology of Aulopyge is unique among cypriniforms. 



Discussion 
Aulopyge relationships and barbin classification 

Aulopyge exhibits a condition well known to cyprinid systematists, namely the possession of 
several unique features (autapomorphies) and few, if any, recognisable synapomorphies with 
other cyprinid taxa. Too few published comparative anatomical data exist for barbelled carps 
(Cyprininae) and the comparisons made during this study are of limited taxonomic scope. 
However, some information has emerged which may signpost useful characters for determining 
subgroups amongst barbins. The phylogenetic position of Aulopyge can only be discussed in the 
context of these wider issues. 

It was stated in the Introduction that Aulopyge is a member of the Cyprininae. This subfamily 
was one of the divisions recognised by Howes (1981) on the basis of: 



CYPRINID FISH GENUS A ULOPYGE 1 87 

*a maxillary barbel associated with a foramen in the maxillary bone through which the barbel is 
supplied by a branch of the VII facial nerve. 

*a rostrally extended supraethmoid with a laterally convex border. 

At present only two monophyletic assemblages have been identified within the Cyprininae, viz. 
the squaliobarbin group (Howes, 1981) and the labein group (Reid, 1982; 1985). The former is a 
group of three seemingly plesiomorphic genera (Ctenopharyngodon, Mylopharyngodon and 
Squaliobarbus) having a native distribution restricted to China. The labeins are a speciose 
assemblage of c. 16 genera with an Afro- Asian distribution. Aulopyge shares none of those derived 
characters listed by Howes (1981) and Reid (1982; 1985) as defining either group. 

The Cyprininae may be subdivided on the basis of the morphology of the dilatator fossa (Howes, 
1981:15). There are two conditions of the fossa; 1 ) it indents the dorso-lateral cranial surface, or 2) 
it is a foramen in the ventral lamella of the frontal, and in the case of the labeins, the sphenotic as 
well. 

It is assumed from its widespread occurrence in teleosts, and its universal presence in all non- 
barbelled cyprinids (Leuciscinae) and other cyprinoids, that the dorsal cranial dilatator fossa 
represents the pleisiomorphic condition. That the ventral, foraminate dilatator fossa is a derived 
condition is reinforced by its ontogenetic history. 

The development of the foraminate fossa was traced in a series ofBarbus intermedius specimens 
20-55 mm SL. In the smallest specimens the fossa is of the plesiomorphic type, (i.e. dorso-laterally 
placed and indenting the surfaces of the sphenotic and frontal); the dilatator operculi muscle is a 
narrow band-like element. At 29 mm SL there is a lateral process on both the frontal and sphenotic; 
the indentation for the muscle in the frontal has deepened. By 31 mm SL the spheiiotic process has 
curved downward and the frontal lamella is perforated; the anterior part of the dilatator muscle 
runs through the foramen and fibres also originate from its lateral rim and the sphenotic process 
(Figs 1 8a-c). By 55 mm SL the foramen is well-formed and increased in size by medial attrition of 
the frontal lamella. 

In the smallest specimens ofBarbus barbus available (25 mm SL) there is no sign of a foraminate 
dilatator fossa and the condition resembles that in the smallest specimen ofBarbus intermedius. It is 
reasonable to assume that the development of the fossa in this species proceeds along much the 
same course as that described in B. intermedius. In the two closely related species Barbus litamba 
from Lake Malawi and B. mattozi from the Limpopo, the fossa is foraminate only in specimens 
above 103 mm SL, and then only has a small opening. 

Although it may be argued that a foraminate dilatator fossa could have been derived inde- 
pendently within different cyprinine lineages, it will be accepted as a working hypothesis that it is 
the principal synapomorphy for one group of Cyprininae. Since Aulopyge lacks a foraminate 
dilatator fossa it must be included with the squaliobarbins, schizothoracins, several Barbus species 
and other taxa listed in Table 3. Of these, the most likely candidate for sister-group relationship to 
Aulopyge is the schizothoracin assemblage. Some schizothoracin genera lack scales, possess a 
narrow ethmoid, serrated last unbranched dorsal fin ray and have a well-developed ventral facet on 
the lateral ethmoid, all derived characters shared with Aulopyge. However, these characters are 
mosaically distributed amongst schizothoracin species, no one taxon possessing all together, and 
so a relationship between Aulopyge would involve only certain species, thus making the schizo- 
thoracins a paraphyletic group. Previous authors, in recognising the subfamily Schizothoracinae 
sensu Berg, 1912, have tacitly assumed monophyly. Such an assumption is based on the possession 
by all included taxa of 'tile' scales, i.e. a row of regular, oblong scales at the base of the anal fin. This 
synapomorphy is supported by another, namely the presence of a bony strut extending from the 
parasphenoid to contact the prootics in the midline and thus dividing the posterior myodome. As 
such, this feature resembles the basisphenoid present in other teleosts, but which is absent in 
ostariophysans. These two characters indicate the monophyly of the schizothoracins and as such 
exclude Aulopyge, which lacks both of them. 

Aulopyge also shares the character of well-developed lateral ethmoid and entopterygoid facets 
with some species of Barbus. This character distribution immediately raises the question; what is 
meant by Barbus] 

At the present time Barbus includes c. 800 nominal species distributed in Eurasia and Africa, 



188 



G. J. HOWES 





Fig. 18 Barbus intermedius intermedius, ontogenetic development of dilatator fossa; (a) at 21 -5 mm; (b) 
at 25 mm; (c) at 3 1 mm SL. Dashed line indicates position of dilatator operculi muscle; all in ventro- 
lateral view. Scales = 0-5 mm. 



many of which, even to the non-specialist, bear scant resemblance to the type species of the genus, 
the European Barbus barbus (Linn.). Some authors have opted to recognise separate genera, (e.g. 
Puntius, Tor) for Indian and South East Asian species, a solution which does little to elucidate 
relationships since these 'genera' are not defined on derived characters. The definition of Barbus 
can only be approached through an adequate anatomical comparison of the Eurasian and African 
species. 

Comparisons and character analyses made during this study have demonstrated that a 'group' 
including the type species Barbus barbus and other Eurasian species can be defined on a suite of five 
characters: 

1) shield-shaped supraethmoid (Fig. 19a) 

2) oblong lachrymal with ventral sensory canal (Fig. lOb) 

3) enlarged lateral ethmoid facet articulating with a well-developed entopterygoid facet (Fig. 5a) 

4) 13-15 pre-dorsal vertebrae 



CYPRINID FISH GENUS AULOPYGE 
Table 3 Distribution of the dilatator fossa morphotypes amongst examined Cyprininae. 



189 



Foramen present: 

Single foramen 

Acrossocheilus 

Barbichthys 

Barbus (part; see Table 4) 

Capoeta 

Carassius 

Cyprinus 

Probarbus 

Varicorhinus 



Foramen absent: 

Ageniogarra 

Aulopyge 

Barbus (part; see Table 4) 

Cyprinion 

Mystacoleucus 

Onychostoma 

Prolabeo 



Double foramen 

Catla 

Cirrhina 

Crossocheilus 

Garra 

Labeo 

Labiobarbus 

Lobocheilos 

Osteocheilus 

Semilabeo 

Tylognathus (sensu Reid, 1985) 

Typhlogarra 



and in the squaliobarbins 

Squaliobarbus 

Mylopharyngodon 




Fig. 19 Ethmoid region in dorsal view of, (a) Barbus bar bus; (b) B. altianalis altianalis; (c) B. leonensis 
(scale bar = 0-5 mm); (d) B. serra; (e) Tor putitora. 



190 G.J.HOWES 

Table 4 Condition of the dilatator fossa in 80 species of 'Barbus\ 



Foramen present 

African species: altianalis (all subspecies), andrewi, biscarensis, callensis, camp I acanthus, fritschi, guirali, 

intermedius (all subspecies), jacksonijohnstoni, litamba (some, see text), macrolepis, marequensis (all morphs), 

natalensis, nigrodor sails, oxyrhynchus,progenys, reinii, rothschildi, serra, setivemensis, somerini, trimaculatus , 

tropidolepis. 

Eurasian species: barbus, barbulus, bocagei, canis, comiza, douronensis, graellsii, grahami, hexagonolepis, 

longiceps, meridionalis , nasus, plebejus, sharpeyi, tambroides, tor, xanthopterus. 

Foramen absent 

African species: ablabes, amphigramma, argenteus, aspilus aurantiacus, dorsolineatus, eutaenia, holotaenia, 
hospes, hypsolepis, kersteni, leonensis, lineomaculatus, macrops, mimus, neglectus, neumayeri, paludinosus, 
paytoni, perince, poechi, profundus, radiatus, tenuis, thalamakanensis. 

Asian species: aurilius, bimaculatus, binotatus, burmanicus, chola, collingwoodi, conchonius, filamentosus, 
pentazona, lithopides, orphoides, sarana, sophore, titteya. 



5) low neural complex widely separated from the 4th neural spine (Fig. 1 Ib) 
Of these only characters 1 and 2, because of their restricted distribution, can be treated as 
synapomorphies (character 2 is also shared with Aulopyge; see below). Characters 3-5, when 
viewed in the context of cyprinoid distribution are apparently derived. Their disparity among 
cyprinines, however, does not make them highly corroborated synapomorphies. Nonetheless they 
are congruent with characters 1 and 2. 

If, on the basis of this character suite, Barbus is restricted to only some Eurasian species (see 
Appendix 1) then it remains to be determined how closely related it is to those African and Asian 
species presently included in Barbus, Tor and Puntius. From the distribution of the foraminate 
dilatator fossa (see above) it is clear that African and Asian barbins do not constitute a mono- 
phyletic assemblage. Of 80 African and Asian 'Barbus' species selected at random, virtually 50% 
possess the character (4 1 with, 39 without; Table 4). Also emerging from this analysis is that almost 
none of the 'small' African Barbus examined possess a foraminate fossa. Thus, on the basis of the 
synapomorphic foraminate fossa, some Barbus species are more closely related to labein and other 
cyprinine genera such as Capoeta, Cyprinus, Varicorhinus and Acrossocheilus than to other Barbus 
species. 

Immediate relationships ofAulopyge 

Although it lacks a foraminate dilatator fossa, Aulopyge shares with some Eurasian Barbus species 
(termed from hereon Barbus sensu stricto) well-developed lateral ethmoid and entopterygoid 
articular facets and an oblong lachrymal with a ventral sensory canal (although in Aulopyge the 
canal is not fused with the bone; see p. 1 74. The lack of a foraminate fossa may be interpreted either 
as a loss or as a plesiomorphic condition, in which case the lateral ethmoid-entopterygoid facets 
and oblong lachrymal must be viewed as having been independently derived. Fewer assumptions 
are required to support the 'loss' of the derived dilator fossa in Aulopyge than are demanded by 
other schemes of relationship (Figs 20A-C). Support for 'loss' is that Aulopyge exhibits heter- 
ochrony in the late development and fusion of infraorbital sensory canals and in the absence of 
scales. It may be that the dorso-lateral dilatator fossa is also the retention of an early ontogenetic 
stage (see p. 187). Outright dismissal of independent origin on grounds of parsimony must be 
treated with caution, however, since it is noted that amongst the schizothoracins a lateral ethmoid 
facet is present in some taxa (p. 1 70). Since the schizothoracins are almost certainly a monophyletic 
group within the Cyprininae (see p. 1 87) it follows that this feature has been derived independently 
from that in Aulopyge, Barbus, sensu stricto and other barbins (including Cyprinus). 
Whether Aulopyge is recognised as the sister-group to Barbus sensu stricto, or to Barbus sensu 



CYPRINID FISH GENUS AULOPYGE 



191 




Fig. 20 Three possible schemes of relationship between Aulopyge and other cyprinines. A, the most 
parsimonious, involves the loss of character 2 in Aulopyge. B. involves the loss of characters 3 and 4 in 
'Barbins' and Labeins. C, involves independent derivation of characters 3 and 4 in Aulopyge and 
Barbus sensu stricto. 

Character 1. lateral ethmoid articular facet; 2. foraminate dilatator fossa; 3. expansion of lateral 
ethmoid facet and presence of entopterygoid facet; 4. oblong lachrymal; 5. double-foraminate fossa 
(additional characters defining Labeins are given in Reid, 1985). 'Barbins' include those taxa listed in 
Table 3, under 'foramen present'. 



stricto plus other barbins and labeins cannot be resolved on those characters considered here. 
However, no synapomorphies have been identified that would suggest Aulopyge is more closely 
related to any cyprinine taxon lacking a foraminate dilatator fossa, including the schizothoracins. 
Karaman's (1971) hypothesis of an intermediate taxonomic position for Aulopyge between 
'barbine' and 'schizothoracine' subgroups (see above, p. 165) is not supported by this study. 
Schizothoracins do share with Barbus sensu stricto high total and pre-dorsal numbers of vertebrae 
(Table 2), but the polarity of this character is uncertain (see p. 182) and if treated as a synapomor- 
phy in a scheme of relationship involving Aulopyge, Barbus and 'other cyprinines' it is incongruent 
with other synapomorphies. 



192 G.J.HOWES 

I have found no evidence in support of Arai's (1982) contention that Aulopyge possesses some 
gobionine characters. 

From the data analysed two hypotheses are available: 

*Aulopye is a derived member of the Barbus sensu stricto lineage, with specialisation through 
reduction and 'loss' (Fig. 20A) 
* Aulopyge is a member of the 'stem-group' of Eurasian plus African barbins (Figs 20B & C). 



Acknowledgements 

I am most grateful to my colleagues Keith Banister, Humphry Greenwood, Alwyne Wheeler and Peter 
Whitehead for their critical comments on the manuscript, and for their many enlivened discussions over the 
years on the taxonomy of Barbus. To Gordon Reid of the Horniman Museum, I am particularly indebted for 
his critical refereeing, his sound advice and continuing encouragement. Paul Skelton of the J. L. B. Smith 
Institute of Ichthyology has my gratitude for providing specimens and so much stimulating discussion. 

My sincerest thanks are due to Primoz Zupancic of Dolsko, Yugoslavia, for obtaining the specimens of 
Aulopyge upon which most of this work is based, and for providing me with photographs and information on 
their habitat; and to Keith Easton of the Severn Trent Water Authority for providing juvenile specimens of 
Barbus barbus. 

Special thanks go to my colleague Mandy Holloway for preparing the radiographs. 

After this paper had been submitted for publication, Dr Friedhelm Krupp of the Senckenberg Museum 
generously provided me with additional information which has been added to the text. 



References 

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1'Afrique du Nord. Arquivos do Museu Bocage 2 a ser., 3(10): 319-330. 
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Bulletin du Museum national d'Histoire naturelle, Paris, 4e ser., 3, Sect. A. No. 1: 277-307. 

1983. Re-examination of the types of Barbus haasi Mertens 1924. Senckenbergiana Biologica 63(1-2): 



33-38. 
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pumila, with notes on Eurasian cyprinids and their karyotypes. Bulletin National Science Museum, Tokyo 

(Zool.) 8(3): 133-1 52. 
Banister, K. E. 1980. The fishes of the Tigris and Euphrates rivers. In: Euphrates and Tigris, Mesopotamian 

ecology and destiny. (Eds Rzoska, J., Tailing, J. F., Banister, K. E.). Junk. The Hague-Boston-London: 

95-108. 

Bleeker, P. 1863. Systema Cyprinoideorum revisum. Nederlandsch Tijdschrift voor de Dierkunde 1: 187-218. 
Berg, L. S. 1912. Faune de la Russie: Poissons 3(1): 1-336. St Petersburg. 
Bonaparte, C. L. 1846. Catalogo metodico dei pesci Europei. Atti della Societa Italiana di Scienze Naturali e 

del Museo Civico di Storia Naturals, Milano: 1-95. 
Boulenger, G. A. 191 1 . Catalogue of the fresh-water fishes of Africa in the British Museum (Natural History) 2: 

529pp. 
Chen Xiang-Lin, Yue Pei-Qi & Lin Ren-Duan. 1984. Major groups within the family Cyprinidae and their 

phylogenetic relationships. Acta Zootaxonomica Sinica 9(4): 424440. 
Gosline, W. A. 1978. Unbranched dorsal-fin rays and subfamily classification in the fish family Cyprinidae. 

Occasional papers of the Museum of Zoology, University of Michigan No. 684: 1-21. 
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of the British Museum of Natural History (Zoology) 19(3): 1 19-135. 
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Bulletin of the British Museum of Natural History (Zool.) 34(1): 1-64. 
1979. Notes on the anatomy of Macrochirichthys macrochirus (Valenciennes). 1844 with comments on 

the Cultrinae (Pisces, Cyprinidae). Bulletin of the British Museum of Natural History (Zool.) 36(3): 147-200. 
1981. Anatomy and phylogeny of the Chinese Major Carps Ctenopharyngodon Steind., 1866 and 



Hypophthalmichthys Blkr., 1860 Bulletin of the British Museum of Natural History (Zool.) 41(1): 1-52. 



CYPRINID FISH GENUS A ULOPYGE 1 93 

1982. Anatomy and evolution of the jaws in the semiplotine carps with a review of the genus Cyprinion 
Heckel, 1843 (Teleostei: Cyprinidae). Bulletin of the British Museum of Natural History (Zool.) 42(4): 
299-335. 

1984. Phyletics and biogeography of the aspinine cyprinid fishes. Bulletin of the British Museum of 



Natural History (Zool.) 47(5): 283-303. 
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American Midland Naturalist 56(1): 204-223. 

Jayaram, K. C. 1981. The freshwater fishes of India. A handbook. Calcutta. 475pp. 
Karaman, M. S. 1971. Siisswasserfische der Turkei 8. Revision der Barben Europas, Vorderasiens und 

Nordafrikas. Mitteilungen aus den Hamburgischen Zoologischen Museum undlnstitut. 67: 175-254. 
Lekander, B. 1949. The sensory line system and the canal bones in the head of some ostariophysi. Acta 

Zoological: 1-131. 
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18, Strasbourg. 269pp. 
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and MR AC, Tervuren: 217-343. 
Lindsey, C. C. 1975. Pleomerism, the widespread tendency among related fish species for vertebral number to 

be correlated with maximum body length. Journal of the Fisheries Research Board of Canada 32(12): 

2453-2469. 
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Thesis, University of London. 369pp. 
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Weberian apparatus of the subfamily Gobioninae (Cyprinidae). Acta Zoologica 36(2): 127-158. 
Reid, G. M. 1982. The form, function and phylogentic significance of the vomero-palatine organ in cyprinid 

fishes. Journal of Natural History 16: 497-510. 
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Zoologicae, 6: 1-322. 

Seeley, H. G. 1886. The freshwater fishes of Europe. Cassell, London, Paris, New York and Melbourne. 444pp. 
Skelton, P. H. 1976. Preliminary observations on the relationships of Bar bus species from Cape coastal rivers, 

South Africa (Cypriniformes, Cyprinidae). Zoologica Africana 11(2): 399-41 1. 
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Africa. Ph.D. Thesis, Rhodes University. 417pp. 
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Leuciscus leuciscus (L.). Bulletin Academie Royale de Belgique 5e ser., 63: 469-479. 

Manuscript accepted for publication 1 1 April 1986 



Appendix 1 
The genus Barbus sensu stricto 

Definition and included species 

Banister (1980) placed the majority of Middle-eastern Barbus into two groups the 'European' 
and 'Afro-Indian', which he characterised on overall morphology, scale type and serrated or 
smooth last dorsal fin ray. Banister stated that both groups might be monopyletic and so tacitly 
restricted Barbus to the 'European' group. 

Leveque & Daget (1984) stated that 'Strictly speaking the generic name Barbus shall be restricted 
to European and some north- African species'. These authors' remarks are supported by this study 
and reference has been made in the text to Barbus sensu stricto. Only a thorough comparative 
anatomical study of 'barbins' will provide a strict diagnosis (based on synapomorphies) of Barbus. 
The definition of Barbus used here is based on the characters analysed above and forms a working 



194 G.J.HOWES 

hypothesis for a more critical evaluation. Those taxa not embraced by this definition are referred to 
as 'Barbus' or Barbus sensu lato; in the case of the Asian species, the generic names Puntius and Tor 
are already widely used (see for example, Jayaram, 1981). For African 'Barbus' several generic 
names are available (see synonymy in Leveque & Daget, 1984). 

Barbus sensu stricto is defined on the basis of its members having a total vertebral count of 40-48 
of which 13-15 are pre-dorsal vertebrae; a well-developed, centrally to anteriorly situated ventral 
lateral ethmoid facet articulating with a well-developed anterodorsal entopterygoid facet; a 
'shield'-shaped supraethmoid with (usually) a prominent rostral process (Fig. 19a); neural com- 
plex low with a deeply indented anterior border, its posterior border (usually) well-separated from 
the 4th neural spine which is at least half the height of the neural complex; lachrymal elongate, 
often oblong with tapered anterior tip and sometimes an indented posteroventral border, sensory 
canal running through the lower half of the bone; 49-90 scales in the lateral line, (cf. 20-55 in 
African and Asian 'Barbus'). 

Almaca (1981) distinguished three groups of Eurasian Barbus on the basis of lateral line scale 
counts but he pointed out the lability of this character due to influences of temperature and 
latitudinal variation (see similar remarks under 'vertebrae', p. 182). 

The following species are considered to constitute Barbus sensu stricto: 
Barbus albanicus Steindachner, 1870 (including B. graecus (Steindachner, 1895)) 
Distribution: Albania-Greece 

Barbus barbus (Linnaeus, 1758), type species of the genus. 

Distribution: Europe (see Almaca, 1981 for detailed distributional data and recognition of 
subspecies) 

Barbus barbulus Heckel, 1846 
Distribution: Tigris 
Barbus bocagei Steindachner, 1 865 
Distribution: Iberia 
Barbus capita (Giildenstadt, 1 773) 
Distribution: Caspian and Aral Sea basins; Amu Darya 
Barbus comiza Steindachner, 1865 
Distribution: Iberia 

This species is included in Barbus on the basis of its possessing a high vertebral number, and a 
typical oblong lachrymal (Fig. lOc). However, it differs from other species in its longer and 
narrower head (see Almaca, 1967; 1972), concave dorsal profile, lower number of pre-dorsal 
vertebrae (1 2, cf. 1 3-15), tall neural complex narrowly separated from the 4th neural spine, and the 
absence of a fleshy overhanging upper lip. In its striking dorsal and lateral head profiles, and 
narrow ethmoid B. comiza greatly resembles Aulopyge. However, no synapomorphies have been 
identified that would suggest these features are anything other than homoplasies. 
Barbus esocinus (Heckel, 1843) 
Distribution: Tigris-Euphrates 
Barbus graellsi (Steindachner, 1 866) 
Distribution: Portugal 
Barbus lacerta Heckel, 1843 
Distribution: Tigris-Euphrates and Qwarq rivers 
Barbus longiceps Valenciennes, 1 842 
Distribution: Jordan River system 
Barbus lorteti Sauvage, 1882 
Distribution: Orontes R. 
Barbus microcephalus Almaca, 1 967 
Distribution: Iberia 

Barbus meridionalis Risso, 1826 (including B. peloponnesius Val., 1842). 
Distribution: NE Spain S. France Yugoslavia Greece 
Barbus nasus Giinther, 1874 
Distribution: Morocco 
Barbus pectoralis Heckel, 1843 



CYPRINID FISH GENUS AULOPYGE 195 

Distribution: Orontes R. 

BarbusplebejusEonaparle, 1839 (including the subspecies recognised by Almac. a, 1981; 1983) 

Distribution: N. Italy-Greece 

Barbus rajanorum Heckel, 1 843 

Distribution: Tigris-Euphrates 

Karaman (1971) considers B. schejch (Heckel, 1843) and B. barbulus Heckel (listed here as a 
separate species), to be synonyms of B. rajanorum. This synonymy is doubtful and the 'rajanorum 
complex' requires a taxonomic reappraisal. In Dr F. Krupp's opinion (pers. com.) B. rajanorum is a 
hybrid between B. pectoralis and Capoeta damascinus. 
Barbus sclateri Giinther, 1 868 
Distribution: Iberia 
Barbus subquincunciatus Giinther, 1 868 
Distribution: Tigris-Euphrates 
Barbus steindachneri Almac. a, 1967 
Distribution: Iberia 
Barbus xanthopterus Heckel, 1843 
Distribution: Tigris-Euphrates 

Although the Middle-eastern species Barbus grypus and B. sharpyeihave relatively high vertebral 
numbers (Table 1), they lack the elongate lachrymal of the other species listed and possess, in 
common with B. canis and B. reinii what is considered to be another derived form of lachrymal in 
which the sensory canal runs along the anterodorsal border (see p. 177). Barbus canis and B. reinii 
both have low vertebral numbers, respectively 38 39 and 37 (totals) and 10 and 10-1 1 predorsal 
elements. Barbus sharpy ei differs from other species of this group in having only 30-31 lateral line 
scales. 



The systematic positions of Barbus brachycephalus Kessler, 1872 and B. mursa (Guldenstadt, 1773) 

The generic placements of these two south Central Asian species (respectively the Aral and 
Caspian Seas and the Kura system) are problematical. Both species although having high vertebral 
counts differ in several ways from Barbus sensu stricto and other species of 'Barbus'. 
Barbus brachycephalus has rather slender barbels, unlike the thick, often papillate barbels of 
'typical' species of the genus, and 7 branched dorsal fin rays, cf. 8 in the majority of Barbus sensu 
stricto and also in Barbus sensu lato. The cranium is broad and flat, lacking the transverse convexity 
of most barbins. There are a total of 47 vertebrae, but only 1 1 are predorsal, cf. 13-15 in Barbus 
sensu stricto. 

Barbus mursa has a deep lachrymal with an anterior branching pattern resembling that of Barbus 
canis and related species discussed above (Fig. lOn). However, it possesses a series of preanal scales 
and a prominent genital papilla more reminiscent of schizothoracins. 



The systematic positions of Barbus andrewi Barnard, 1937 and B. serra Peters, 1864 

These two species are restricted to the South African Western Cape. On the basis of the characters 
given for Barbus sensu stricto both species should be included. Both, however, have a higher pre- 
dorsal vertebral count than other Barbus sensu stricto, viz.: 14-17, cf. 13-15, but, a relatively low 
total vertebral number, viz.: 38-41, cf. 40-47. The supraethmoid has the same 'shield'-shaped 
appearance as in Barbus sensu stricto (Fig. 19d), but the vomer is broader anteriorly and extends 
further dorsally in B. serra and B. andrewi. Again, the lachrymal bones of the two species, while 
having the same overall appearance of that bone in the Eurasian Barbus have a sloped, rather than 
a perpendicular posterior margin and more convex ventral border. Because of these differences I 
am hesitant to include the Cape species in Barbus sensu stricto. According to Skelton (1 980), Barbus 
serra and B. andrewi are sister-taxa, not closely related to any African 'Barbus' he examined. 



196 



G. J. HOWES 



Appendix 2 
Characteristics of the subfamilies Cyprininae and Leuciscinae 



CYPRININAE 
(including 'Schizothoracinae') 

Maxillary barbel present 
Maxillary foramen 

Supraethmoid with medially indented rostral 
process 



4 Articular facets present on lateral ethmoid and 
entopterygoid in some taxa 

5 Single, strong ligament connecting posterior face 
of lateral ethmoid with dorsomedial surface of 
entopterygoid 

6 Lachrymal (1st infraorbital) sometimes elongate 
and oblong 

7 Dilatator operculi muscle sometimes originating 
from ventral surface of the frontal and passing 
through a frontal-sphenotic foramen 

8 Neural complex lacking grooved dorsal surface; 
sometimes close to, or even contacting the cranium 
(supraoccipital) 

9 1 st free supraneural not contacting neural complex 



10 Dorsal fin origin lies above or anterior to that of the 
pelvics (rarely posterior) 

11 1st unbranched dorsal fin ray lies above the 
llth- 18th vertebra 

1 2 Last unbranched dorsal fin ray often serrated 



LEUCISCINAE 

No maxillary barbel 

No maxillary foramen 

No supraethmoid rostral process 

Articular facets absent (except in Tinea) 

Connection between lateral ethmoid via connec- 
tive tissue sheet, sometimes thickened laterally 

Lachrymal never elongate or oblong 



Dilatator operculi muscle confined to cranial 
surface; fossa never foraminate 



Neural complex with grooved dorsal surface, 
never contacting the cranium 



1st free supraneural articulating with neural 
complex 

Dorsal fin origin rarely above or in advance of the 
pelvics 

1st unbranched dorsal fin ray lies above 16th-31st 
vertebra 

Last unbranched dorsal fin ray never serrated 



1 3 Modal number of vertebrae 38^40 (never exceeding Modal number of vertebrae 40-45 (range 33-61 ) 
48) 

Note: The genus Tinea is included here in the Leuciscinae, although possessing a cyprinine feature 
(character 4). Chen et al. (1984) consider Tinea to be the sister-group of the Cyprininae (their 
'Barbines'). Although for the most part, these authors appear to base their hypothesis on differ- 
ences rather than on shared homologies, their cladogram requires serious consideration and offers 
a much needed, testable classification of the Cyprinidae. 






British Museum (Natural History) 

The birds of Mount Nimba, Liberia 

Peter R. Colston & Kai Curry- Li ndahl 



For evolution and speciation of animals Mount Nimba in Liberia, Guinea and the Ivory Coast is 
a key area in Africa representing for biologists what the Abu Simbel site in Egypt signified for 
archaeologists. No less than about 200 species of animals are endemic to Mount Nimba. Yet, this 
mountain massif, entirely located within the rain-forest biome, is rapidly being destroyed by 
human exploitation. 

This book is the first major work on the birds of Mount Nimba and surrounding lowland 
rain-forests. During 20 years (1962-1982) of research at the Nimba Research Laboratory in 
Grassfield (Liberia), located at the foot of Mount Nimba, scientists from three continents have 
studied the birds. In this way Mount Nimba has become the ornithologically most thoroughly 
explored lowland rain-forest area of Africa. 

The book offers a comprehensive synthesis of information on the avifauna of Mount Nimba 
and its ecological setting. During the 20 years period of biological investigations at Nimba this in 
1962 intact area was gradually opened up by man with far-reaching environmental consequences 
for the rain-forest habitats and profound effects on the birds. Therefore, the book provides not 
only a source of reference material on the systematics, physiology, ecology and biology of the 
birds of Mount Nimba and the African rain-forest, but also data on biogeography in the African 
context as well as conservation problems. Also behaviour and migration are discussed. At 
Nimba a number of migrants from Europe and/or Asia meet Afrotropical migratory and 
sedentary birds. 

Professor Kai Curry-Lindahl has served as Chairman of the Nimba Research Laboratory and 
Committee since its inception in 1962. Peter Colston is from the Subdepartment of Ornithology, 
British Museum (Natural History), Tring, and Malcolm Coe is from the Animal Ecology 
Research Group, Department of Zoology, Oxford. 

1 986, 1 29pp. Hardback. 565 00982 6 1 7.50. 






Titles to be published in Volume 52 



Miscellanea 

A revision of the Suctoria (Ciliophora, Kinetofragminophora) 5. The Paracineta 
and Corynophora problem. By Colin R. Curds 

Notes on spiders of the family Salticidae 1. The genera Spartaeus, Mintonia and 
Taraxella. By F. R. Wanless 

Mites of the genus Holoparasitus Oudemans, 1936 (Mesostigmata: Parasitidae) in 
the British Isles. By K. H. Hyatt 

The phylogenetic position of the Yugoslavian cyprinid fish genus Aulopyge Heckel, 
1841, with an appraisal of the genus Barhus Cuvier & Cloquet, 1816 and the 
subfamily Cyprininae. By Gordon J. Howes 

Revision of the genera Acineria, Trimyema, and Trochiliopsis (Protozoa, 
Ciliophora). By H. Augustin, W. Foissner & H. Adam 

The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) with a 
systematic review, a synopsis of Pipistrellus and Eptesicus, and the descriptions of 
a new genus and subgenus. By J. E. Hill & D. L. Harrison 

Notes on some species of the genus Amathia (Bryozoa, Ctenostomata). 

By P. J. Chimonides 



Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 



Bulletin of the 

British Museum (Natural History) 



Revision of the genera Acineria, Trimyema 
and Trochiliopsis (Protozoa, Ciliophora) 



H. Augustin, W. Foissner & H. Adam 



Zoology series Vol 52 No 6 25 June 1987 



The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four 
scientific series. Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and 
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Zool.) 



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ISBN 565 05030 3 

ISSN 0007- 1 498 Zoology series 

Vol52 No. 6 pp 197-224 

British Museum (Natural History) 

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London SW7 5BD Issued 25 June 1987 



(NAT 

26 
Revision of the genera Acineria, Trimyema and 

Trochiliopsis (Protozoa, Ciliophora) 

H. Augustin, W. Foissner & H. Adam 

Zoologisches Institut der Universitat Salzburg, Hellbrunnerstrasse 34, Salzburg, Austria, A-5020 



Conte 




Synopsis . 
Introduction . 
Materials and Methods 
Genus Acineria 
Genus Trimyema 
Genus Trochiliopsis . 
References 



Synopsis 

The genera Acineria, Trimyema and Trochiliopsis are reviewed. The revision is based on an investigation of 
each of the type-species, namely Acineria incurvata Dujardin, Trimyema compressa Lackey, and Trochiliopsis 
opaca Penard, which were found in a sewage-treatment plant. Acineria comprises three species; A. incurvata, 
A. nasuta, and A. uncinata. A. acuta is a synonym of A. incurvata. Trimyema comprises eight species; 
T. alfredkahli, T. claviformis, T. compressa, T. echinometrae, T. kahli, T. marina, T. minuta and T. pleuri- 
spiralis but T. alfredkahli and T. claviformis are perhaps synonyms of T. marina. Trochiliopsis is monotypic 
and new for the fauna of Austria. This genus is apparently closely related to the autochthonous soil ciliate 
Stammeridium kahli. 



Zusammenfassung 

Die Gattungen Acineria, Trimyema und Trochiliopsis werden revidiert. Die Revision basiert auf der 
Untersuchung der Typusarten, namlich Acineria incurvata Dujardin, Trimyema compressa Lackey und 
Trochiliopsis opaca Penard, die in einer Klaranlage gefunden wurden. Die Gattung Acineria umfasst drei 
Arten; A . incurvata, A . nasuta und A . uncinata. A . acuta ist ein Synonym von A . incurvata. Von Trimyema sind 
acht Arten beschrieben; T. alfredkahli, T. claviformis, T. compressa, T. echinometrae, T. kahli, T. marina, T. 
minuta und T. pleurispiralis. T. alfredkahli und T. claviformis sind moglicherweise Synonyme von T. marina. 
Trochiliopsis ist monotypisch und neu fur die Fauna Osterreichs. Diese Gattung ist hochstwahrscheinlich 
nahe verwandt zum autochthonen Boden-Ciliaten Stammeridium kahli. 



Introduction 

Only few activated-sludge ciliates have been characterized by silver-staining techniques which is 
sometimes necessary for their correct identification. To overcome this deficiency, a project to 
redescribe the most frequently occurring species was begun. During these studies the poorly known 
type-species of the genera Acineria, Trimyema, and Trochiliopsis were found. They have been 
reinvestigated using modern techniques which provide a base upon which to revise these genera. 

Supported by the 'Fonds zur Forderung der wissenschaftlichen Forschung, Projekt Nr. P 5889'. 



Bull. Br. Mus. nat. Hist. (Zool.) 52(6): 197-224 Issued 25 June 1987 

197 



1 98 H. AUGUSTIN, W. FOISSNER & H. ADAM 

Materials and Methods 

Acineria incurvata, Trimyema compressa, and Trochiliopsis opaca were obtained from activated 
sludge of the sewage-treatment plant at Aspach, Upper Austria. 

Small samples of activated sludge were placed in glass petri-dishes where they remained without 
additional aeration. In such cultures a surprising succession and enrichment of ciliates often 
occurred. Acineria incurvata could also be cultured in tap water enriched with a crushed wheat 
grain which supported the growth of many small prey ciliates (Dexiotricha, Uronemd). 

The infraciliature was revealed with a protargol silver-staining method (Foissner, 1982). The 
silverline system was studied in specimens impregnated by a modified 'dry' silver-impregnation 
technique (Foissner, 1976). The oral structures of Trimyema compressa were impregnated by the 
pyridinated silver carbonate method of Fernandez-Galiano (1976) as improved by Augustin et al. 
(1984). 

For scanning electron microscopy Acineria cells were fixed for 10 minutes in Parducz's solution 
(2% OsO 4 and concentrated Hg-sublimate solution, 4:1), rinsed in 0-05 M sodium cacodylate 
buffered at pH 6-3, dehydrated in an isopropyl alcohol series (60%, 70%, 80%, 90%, 100%, 100%, 
five minutes each) and put into a mixture of isopropyl alcohol (100%) and frigen 1 1 (2 : 1, 1 : 1, 
1 : 2, five minutes each). Finally, cells were transferred into pure frigen 1 1 and critical point dried, 
using frigen 13. Specimens were gold-sputtered three times for six minutes each. 

Each species was drawn from life as well as from impregnated specimens using a camera lucida 
for the latter. The drawings are only slightly diagrammatic. All statistical procedures follow 
methods described in Sokal & Rohlf (1981). 

Genus ACINERIA Dujardin, 1841 

DIAGNOSIS. Amphileptidae Biitschli, 1889 with (1) compressed oral slit anteriorly rolled up and 
overlapping to the left side forming (together with the anterior dorsal margin) an oblique spoon- 
like excavation, (2) three perioral kinetics (one left and two right of the cytostome, (3) somatic 
kinetics on the right side successively shortened along the cytostome, (4) oral slit measuring less 
than half of body length being located at the convex side of the tapering anterior. Freshwater and 
marine, prefers polysaprobic conditions. 

TYPE-SPECIES. Acineria incurvata Dujardin, 1841 

REMARKS. Acineria was mentioned for the first time by Dujardin (1840) in the family 'Trichodiens' 
but without any valid characterization. In 1 84 1 he gave a rather vague diagnosis of the genus and of 
two species. Maupas (1883) criticized the unsatisfactory diagnosis and gave a better description 
of Acineria incurvata even noting the overlapping anterior end which is the main character of 
Acineria; nevertheless, he did not include this character in the diagnosis. How Maupas (1883) 
arrived at the conclusion that his species was the same as that described by Dujardin remains, 
however, inexplicable. It was only Kahl (1926) who used the real character of the overlapping 
dorsal end of the mouth to the left side to distinguish Acineria from the most closely related genus 
Litonotus. But there is no indication in the infraciliature that the dorsal margin and the left side 
coalesce as supposed by Kahl (1926, 1931). Thus, Kahl's interpretation that a part of the ciliated 
right side of the genus Litonotus has shifted over to the left side in Acineria is not supported by our 
investigations. We consider the rolled up anterior part of the mouth to be the reason for the 
anterior overlapping of the dorsal margin. The occurrence of somatic kinetics on the left side, as 
stressed by Kahl (1926) is a weak distinctive character because this happens also, more or less 
pronounced, in the genera Litonotus and Amphileptus (Foissner, 1984). 

Key to the species 

la Single spherical macronucleus A.nasuta 

Ib Macronucleus in two parts with a single micronucleus between them 2 

2a Cytostome restricted to the rolled up anterior pole, right side with 3 somatic kinetics, left side 

unciliated A.uncinata 

2b Cytostome about one third of body length, 10-12 normally ciliated somatic kinetics A. incurvata 



THE GENERA ACINERIA, TRIMYEMA AND TROCHILIOPSIS 199 

Descriptions of species 
Acineria incurvata Dujardin, 1841 

?Trachelius anaticula Ehrenberg, 1 833 

Acineria acuta Dujardin, 1841 

?Amphileptus anaticula Claparede & Lachmann, 1859 

Lionotus reversus Kahl, 1926 

Amphileptus incurvatus Lepsi, \926a 

Litonotus lamella Fryd-Versavel et al., 1975 

NEOTYPE-SPECIMENS. Slide (protargol silver impregnated) of neotype-specimens has been deposited 
in the British Museum (Natural History) in London, reference number 1986:5:30:1 . 

REDESCRIPTION (Figs 1-3, 12-58, Table 1). Type species of the genus. Freshwater and marine. 
About 45-200 urn (Dujardin, 1841; Maupas, 1883; Kahl, 1926, 1928, 1931, 1933; Horvath & 
Kuhn, 1941; Bick, 1972; Foissner, 1977/78). Abnormal, giant individuals up to 500 jam showing 
most organelles duplicated observed by Foissner (1977/78) and probably by Lepsi (1965) (Figs 
28-30). Body oblong, slightly contractile, laterally compressed, rounded posteriorly, narrowing 
anteriorly to a blunt point. Rather variable in shape (slender to wide and plump) depending on 
nutritional condition (Figs 3 1-34). Ventral side more or less convex, dorsal side straight or concave 
in the anterior, convex in the posterior region. Excavated region conspicuous, shining brightly. 
Anterior-most dorsal top somewhat refractive, due to the rolled up oral slit. Macronucleus in two 
spherical to ovoid parts with a single micronucleus between them. 1-3 micronuclei according to 
Maupas (1883). Macronuclear parts fuse during bipartition (Horvath & Kuhn, 1941) (Figs 38-42) 
and divide in the later fission stages (Kahl, 1926). Single contractile vacuole at the posterior pole, 
diameter about 7 um, with 5-8 pores on the right lateral side (Horvath & Kuhn, 1941) (Fig. 43) 
which could not be seen in our slides. Cytoproct terminal, a slightly laterally located slit (Maupas, 
1883; Kahl, 1926). Pellicle soft, flexible, with longitudinal furrows in which the cilia and bristles 
originate. Furrows disappear in well-fed individuals. Extrusomes straight to slightly fusiform 
(arrow-shaped according to Foissner, 1977/78), thin, about 4 um long (2 urn according to Horvath 
& Kuhn, 1941), located along the cytostome, a small accumulation of them in the ventral side of the 
posterior end and even a few scattered throughout the body (Figs 48, 49). Cytoplasm of 
normally-fed specimens rather clear, containing some small colourless spheres. Carnivorous, feeds 
on small hymenostome ciliates, e.g. Colpidium, Cyclidium, Glaucoma, Pseudocohnilembus, Loxo- 
cephalus, Uronema (Maupas, 1883; Lepsi, 1926a; Kahl, 1926, 1931; Buck, 1961; Struhal, 1969). 
Starved individuals feed even on 'cysts' ofEuglena viridis (Horvath & Kuhn, 1941) and perhaps on 
bacteria (Lepsi, 1926a). Ingestion vacuoles rather large, dividing quickly into smaller food 
vacuoles (Horvath & Kuhn, 1941). Movement moderately quick, gliding on the bottom of the 
petri-dish or swimming in rotation along its longitudinal axis. Bipartition by transverse fission 
(Lepsi, 1926a; Horvath & Kuhn, 1941) (Figs 38-42). Opisthe almost spherical when it separates 
from the proter (Kahl, 1926; Horvath & Kuhn, 1941) (Fig. 40). Very small degenerative forms tend 
to conjugate; during this process the mouth of an individual fuses with the back of another (Kahl, 
1926) (Fig. 20). Encystment frequently occurring when food is depleted (Horvath & Kuhn, 1941). 
Endocyst forms within an hour, later the macronuclear parts fuse to a worm-shaped product. Wall 
of ectocyst without visible structure. Cysts surrounded by some material which sticks them to the 
bottom of the culture dishes or to the bacterial film on the surface of the culture medium (Horvath 
&Kuhn, 1941) (Fig. 35). 

Three different types of cilia: (1) normal cilia, about 10 um, (2) short bristles, about 0-5-1-0 um, 
(3) club-shaped bristles, up to 2-0 um. Eleven longitudinal kinetics with cilia type 1, about 8-9 of 
them on the right and about 3 on the left side. This is in accordance with the numbers given by Kahl 
( 1 926), Horvath & Kuhn ( 1 94 1 ), and Fryd-Versavel et al. ( 1 975). In addition to the normal somatic 
kinetics the following are found on the more differentiated left side: (1) a single kinety with cilia 
type 2 located to the left of the brosse kinety and often extending only to the middle of the body, its 
posterior basal bodies less closely spaced, (2) one brosse row of obliquely arranged, paired bristles 
(cilia type 3) being posteriorly continued by a row of unciliated kinetosomes (or by kinetosomes 



200 



H. AUGUSTIN, W. FOISSNER & H. ADAM 




THE GENERA ACINERIA, TRIMYEMA AND TROCHILIOPSIS 201 

with very short bristles only), (3) one kinety consisting apically of 2-3 cilia of type 3 (probably 
constituting a rudimentary brosse row) and being continued by a few unciliated kinetosomes 
(about 5 in the anterior third and about 3 kinetosomes in the middle of the body). Kahl (1926, 
1931) described the brosse as being built up of 3 rows of bristles (Fig. 21). Foissner (1977/78) 
observed only a file-shaped structure there, most probably suggested by the single row of paired 
brosse-bristles. 

Cytostome more or less curved, anteriorly overlapping to the left side but not to the right as 
described by Lepsi (\926a, b, 1928). Perioral kinety 1 left of cytostome, with paired basal bodies 
along the mouth, however, only the anterior basal body each bearing cilia of type 2. Perioral kinety 
2 and 3 to the right of the oral slit showing closely spaced basal bodies and constituting the so-called 
'mane', a conspicuous compact ciliature. Perioral kinety 2 with paired basal bodies along the oral 
slit, the anterior basal body bears a cilium of type 1 . This kinety appears unciliated post-orally. 
Perioral kinety 3 with single basal bodies but ciliated along the whole body with cilia type 1. 
Horvath & Kuhn (1941) misinterpreted the perioral kinetics 2 and 3 as left and right perioral 
kinetics. Their drawing, however, shows the correct situation, that is to say also perioral kinety 1 
(Figs 43, 44). Fryd-Versavel et al. (1975) overlooked the perioral kinety 3 (Figs 45-47). 

The silverline system is a linearly orientated fine-meshed lattice (Foissner, 1977/78) (Fig. 50a). 

OCCURRENCE AND ECOLOGY. Dujardin (1841) found this species in a 20-day-old infusion of material 
from the Mediterranean Sea. Later it was recorded from the brackish waters of Oldesloe and Kiel 
(Kahl, 1928, 1933), from the Roumanian littoral of the Black Sea (Lepsi, \926a,b, 1928;Tucolesco, 
1962a) and from the periphyton of brackish and marine waters of Konigshafen near List (Sylt, 
Germany) (Kiisters, 1974). 

Some authors mentioned also terrestrial habitats (Radu & Tomescu, 1972; Tomescu, 1978), but 
a reliable record is not available (Foissner, 1987). The drawing made by Stella (1948), who claimed 
to have found Acineria incurvata in a pine forest, indicates that it was (probably) a member of the 
genus Spathidium (Fig. 22). 

Acineria incurvata has been frequently found in strongly saprobic freshwater habitats, such as 
different sewage-loaded watercourses (Horvath & Kuhn, 1941; Buck, 1961; Bick, 1972; Madoni & 
Ghetti, 1977; Foissner, 1977/78), in Sphaerotilus tufts (Vasicek, 1964; Struhal, 1969), on the 
bottom of the river Elbe upstream from Hamburg (Grimm, 1968), in a cesspool (Kahl, 1926), and 
in sewage-treatment plants (trickling filters in good working order, aeration tanks) (Buck, 1961; 
Weninger, 1971; Madoni, 1981). Fryd-Versavel et al. (1975) found their 'Litonotus lamella' in a 
pond in the year 1962. Sramek-Husek (1956, 1958) noted it as a true member of the 'Colpidietum 
colpodae'. Weninger (1971) found a decreasing abundance when nitrate or ammonium was added 
to sewage, whereas phosphate strongly increased its number. 

The above data suggest that Acineria incurvata is a widely distributed poly saprobic eury hyaline 
indicator species with a rather high tolerance of lack of oxygen and high concentrations of NH 4. 



Figs 1-23 Acineria. 

Figs 1, 2 Acineria incurvata after Dujardin (1841). 

Fig. 3 Acineria acuta after Dujardin (1841). 

Fig. 4 Acineria nasuta after Lepsi ( 1 962). 

Figs 5-11 Acineria uncinata after Tucolesco (\962a). 5 Anterior pole. 6 Posterior pole, 7, 8 Right and 
left side. 9 Mouth and anterior pole overlapping towards the left side. 10 Ventral view. 11 Dorsal view. 

Figs 12-23 Acineria incurvata. 12-15 After Maupas (1883). 12 An individual swallowing an Uronema. 
13 Right side (Maupas called it dorsal view). 14 Left side (Maupas called it ventral view). 15 Pellicle. 16 
After Kahl (1931), left side. 17-21 After Kahl (1926). 17, 18 Left side of different individuals. 19 Right 
side. 20 Conjugants. 21 Left anterior region. 22 After Stella (1948), probably a Spathidium. 23 After 
Buck (1961). 



202 H. AUGUSTIN, W. FOISSNER & H. ADAM 

Table 1 Biometrical characterization of Acineria incurvata 



Character 1 



M SD SE CV Min Max n 



Body, length 
Body, width 

Number of macronucleus parts 
Macronucleus part, length 
Macronucleus part, width 
Number of micronuclei 
Micronucleus, length 
Micronucleus, width 
Cytostome, length (measured as chord) 
Distance from apex to posterior end of brosse 
Number of brosse-bristles 
Brosse-bristles, maximal length 
Number of left perioral kinetics 
Number of right perioral kinetics 
Number of normally ciliated kinetics (cilia type 1), 
perioral kinetics excluded 



56-25 54-5 7-50 1-67 13-3 46-0 75-0 20 
15-50 16-0 2-01 0-45 13-0 12-0 19-0 20 



2-00 
9-65 
7-35 
1-00 
2-42 
2-12 
22-55 
16-90 
41-20 
1-72 
1-00 
2-00 


2-0 
10-0 
7-5 
1-0 
2-2 
2-0 
22-0 
19-5 
40-0 
1-8 
1-0 
2-0 


0-00 
1-60 
0-83 
0-00 
0-66 
0-47 
3-50 
2-66 
4-18 
0-24 
0-00 
0-00 


0-00 
0-36 
0-18 
0-00 
0-15 
0-11 
0-78 
0-60 
0-93 
0-05 
0-00 
0-00 


0-0 
16-6 
13-3 
0-0 
27-1 
22-3 
15-5 
13-6 
10-1 
14-3 
0-0 
0-0 


2-0 
7-0 
6-0 
1-0 
1-8 
1-6 
15-0 
14-0 
34-0 
1-2 
1-0 
2-0 


2-0 
13-0 
9-0 
1-0 
4-0 
3-6 
28-0 
25-0 
48-0 
2-0 
1-0 
2-0 


20 
20 
20 
20 
20 
20 
20 
20 
20 
20 
20 
20 



10-85 11-0 0-59 0-13 5-4 10-0 12-0 20 



'All data are based on protargol silver impregnated specimens. All measurements in |im. Legend: x, mean; M, median; SD, 
standard deviation; SE, standard error of mean; CV, coefficient of variation in %; Min, minimum; Max, maximum; n, 
sample size. 

REMARKS. Trachelius anaticula Ehrenberg, 1833 is an older but unreliable synonym of this species. 
Acineria acuta Dujardin, 1841, which was observed in the water of a wheel-track in 1838, has been 
very insufficiently described and therefore cannot be discriminated from Acineria incurvata. Thus, 
Acineria acuta is here treated as synonym. Amphileptus anaticula perhaps is a synonym, too, but 
the figure given by Claparede & Lachmann (1859) shows an unidentifiable individual with a 
voluminous ingestion vacuole. The synonym Lionotus reversus Kahl, 1926 results par lapsus, since 
Kahl mentioned in a footnote that he had found Maupas' good description of Acineria incurvata 
just after having finished the manuscript. The synonym Litonotus lamella results from an obvious 
misidentification by Fryd-Versavel et al. (1975). 

Acineria nasuta Lepsi, 1962 

DIAGNOSIS (Fig. 4). Marine. About 90-100 urn long, rather wide. Only one single macronucleus. 
Pellicle with 5-6 distinct stripes. Postapical, to the right of the so-called 'nose' a peculiar line 
(perhaps the mouth) which is said to be characteristic of this species. 

OCCURRENCE AND ECOLOGY. Only a few individuals were found in a raw culture of putrefying 
marine algae. In the same culture Holophrya torquabilis occurred in large numbers, probably 
serving as food for Acineria. 



Figs 24-^47 Acineria incurvata. 24-30 After Lepsi (1965). 24 Normal aspect. 25 Trachelius-\ike form. 
26 Slender form, resembling Spathidium. 27 Degenerated individual resembling Litonotus. 28-30 
Abnormal, degenerated forms. 31^14 After Horvath & Kuhn (1941). 31-34 Outlines of well-fed and 
starved specimens. 35 Cyst. 36 Left side, with extrusomes along the cytostome. 37 Right side, location 
of contractile vacuole and of kinetics. 38-42 Bipartition. 43, 44 Infraciliature (right and left side) 
revealed by Bresslau's opalblue-technique. 45-47 After Fryd-Versavel et al. (1975), misidentified as 
Litonotus lamella. 45, 46 Infraciliature of left and right side. 47 Diagram of different types of cilia and 
bristles in the anterior region. 



THE GENERA ACINERIA, TRIMYEMA AND TROCHILIOPSIS 



203 




46 



47 



204 



H. AUGUSTIN, W. FOISSNER & H. ADAM 




49 







Figs 56-58 Acineria incurvata, scanning electron micrographs. 56 Total view of left side. 57 Anterior 
part with dorsal oral region rolled up forming a spoon-like excavation. Note the club-shaped brosse- 
bristles, the short bristles of the perioral kinety 1 , and the long cilia of perioral kinetics 2 and 3 (arrows). 
58 Detail of anterior third with brosse-bristles, short bristles and normal cilia (arrows). 



Figs 48-55 Acineria incurvata, originals. 48 Left side from life and according to scanning electron 
microscopic observations, scale = 20 um. 49 Extrusome, length about 4 um. 50 Reconstructed cross- 
sections in different regions of body. 50a Silverline system in the oral region, dry silvered, after 
Foissner (1977/78). 51 Right side, infraciliature of a protargol silver stained specimen. P2, P3, perioral 
kinetics 2 and 3. 52 Left ventro-lateral view of a protargol silver stained specimen with different types 
of cilia and bristles according to SEM-observations. 53, 54 Infraciliature of the left ventro-lateral and 
the right dorso-lateral side of a protargol silver impregnated specimen. Pl-3, perioral kinetics 1-3; Br, 
srale = 30 um 55 Ventral view. 



206 H. AUGUSTIN, W. FOISSNER & H. ADAM 

REMARKS. Lepsi (1962) assumed that this species, which has remained unmentioned since 
original description, could be a form of A. incurvata and mentioned some relationship with the 
genera Chilophrya and Plagiocampa. His figure and description are so incomplete that it is at 
present impossible to find any reliable affinity. The single macronucleus suggests that it is not an 
Amphileptidae, although he could have observed a dividing stage with fused macronucleus. 

Acineria uncinata Tucolesco, 19620 

DIAGNOSIS (Figs 5-1 1). Brackish and freshwater. About 35-55 urn. Body lanceolate without lateral 
edge. Anterior pole overlapping towards the left side. Two spherical macronuclei showing a clearer 
zone at their central region. Sometimes a single, elongated, tapered nucleus. Contractile vacuole 
terminal, often surrounded by a group of smaller vacuoles. Cytostome a straight and short slit 
restricted to the rolled up anterior pole. Can therefore feed only on small prey (flagellates). Three 
somatic kinetics on the right side with 20-22 cilia each. Cilia at the ventral margin of the anterior 
third transformed to regularly curved crotchets. 

OCCURRENCE AND ECOLOGY. This species was found in summer 1954 in a small dirty brackish 
puddle near Lake Tekirghiol and in mesosaprobic freshwaters of Bucarest. 

REMARKS. Tucolesco (1962a) separated this species from A. incurvata by the non-overlapping 
post-oral dorsal margin. However, in A incurvata the situation is rather similar (page 199). Thus, 
we propose the following characters for discrimination from A. incurvata: the presence of only 
three somatic kinetics on the right side, the (probably) unciliated left side, and the short oral slit 
being restricted to the anterior pole. Unmentioned since description. Note after proof reading: 
This is a valid species which we rediscovered recently! Redescription is in preparation. 



Genus TR1MYEMA Lackey, 1925 
Sciadostoma Kahl, 1926 

DIAGNOSIS. Trimyemidae Kahl, 1933 (syn. Sciadostomatidae Kahl, 1926) with vestibulum and 
cytostome near apical end. Vestibular ciliature consisting of three rows of cilia, two rather long 
ones arranged approximately in a semicircle at the left margin of the vestibulum and an inner rather 
short third row located near the cytostome at the posterior left of the vestibulum. Somatic ciliature 
in longitudinal kineties but arranged in a way that a more or less wide band of oblique spirals is 
formed. Prominent caudal cilium. Body small, mostly tapered at both ends. Free-living and 
endocommensally, freshwater and marine, polysaprobic. 

TYPE-SPECIES. Trimyema compressa Lackey, 1925 

REMARKS. There is much confusion about the exact orientation of the cell: dorsal, lateral, and 
ventral sides are often mixed up in descriptions. In addition some authors have given incorrect 
figures focusing the microscope on the lower surface of their specimens. Thus, they attained 
inverted figures (see explanations to figures). Most species of the genus Trimyema are only super- 
ficially described. The oral structures are known exactly only of T. compressa (Figs 83, 107) and 



Figs 59-76 Trimyema compressa. 59 After Lackey ( 1 925) (inverted). 60 After Wang & Nie ( 1 935), left 
lateral view. 61-65 After Liebmann (1936). 61 Left ventro-lateral view. 62-65 Defecation by the aid of 
the caudal cilium. 66-69 After Kahl (1926). 66 Left lateral view. 67 Oral region during progressed 
bipartition. 68 Ventral side. 69 Bipartition. 70 After Kahl (1931), constant marine form, rather similar 
to T. claviformis described later. 71 After Kahl (1933), left lateral view. 72 After Pennak (1953) 
(inverted). 73 After Bick ( 1 972), left lateral view. 74 After Sladecek ( 1 972), left lateral view. 75, 76 After 
Schmall (1976), infraciliature of protargol silver stained specimens (inverted), ventral and dorsal view 
(Schmall called it dorsal and ventral view). 



THE GENERA ACINERIA, TRIMYEMA AND TROCHILIOPSIS 



207 




76 



208 H. AUGUSTIN, W. FOISSNER & H. ADAM 

partly of T. pleurispiralis (Fig. 96) and T. echinometrae (Fig. 93). From the descriptions and our 
investigations we deduced the basic structure of the oral apparatus as described above (compare 
Fig. 1 07). Faure-Fremiet ( 1 962) and Borror ( 1 972) obviously overlooked the short third vestibular 
kinety. Borror (1972) described only an inner and an outer 'polykinety'. Detcheva el al. (1981), 
however, showed in T. compressa electronmicroscopically that, despite their polykinetal appear- 
ance, the vestibular ciliary systems are not separate polykineties but are the anterior parts of the 
somatic kinetics that are preceded by parasomal sacs and retain the same fibrillar systems as the 
somatic kinetosomes. Jankowski (\964a, b) gave no evidence for his statement that there were four 
vestibular kinetics in T. compressa. 

Encystment is unknown in this genus. Czapik (19750) noted that even starved specimens (of T. 
compressa) die without forming cysts. Morphogenesis has not yet been exactly studied. However, 
the oral apparatus is supposed to reduce before cell division, because during division both proter 
and opisthe show the same state of development of the oral apparatus (Kahl, 1926) (Figs 67, 69). 

The silverline system has been demonstrated only in T. compressa (Klein, 1930; Faure-Fremiet, 
1962; Jankowski, \964a, b; Czapik, 19750). Klein (1930) gave the description that best agrees with 
our observations (Figs 106, 1 10). But he did not draw the transverse silver lines connecting the 
longitudinal lines in the region of the ciliary spirals. The granules located at and in the silverlines 
(Fig. 1 10) have been said to be mucocysts ('Relationskorner') or rudimentary basal bodies (Klein, 
1930). However, the electronmicroscopic investigation shows only mucocysts (Detcheva et al., 
1981). 

The exact taxonomic position of the genus is still unclear. Kahl (1926) created a new family for 
the rather special helical ciliature. This author, Corliss (1979), and Curds (1982) included the 
family in the order Trichostomatida Biitschli, 1889. Faure-Fremiet (1962) noted that the family 
Trimyemidae indeed presents one of numerous possibilities existing in the order to use the anterior- 
most somatic kinetics for building up a vestibular ciliature. In addition, he indicated possible 
affinities of Trimyema with Mycterothrix and Maryna, which are now 'good' colpodids (Foissner, 
1985a). Jankowski (1980) erected the new order Trimyemida (incertae sedis) giving no reasons for 
this decision. On the contrary, Detcheva et al. (1981) stated that Trimyema is a member of the 
Vestibulifera and that the Trimyemidae show the same general type of vestibular architecture as 
the Plagiopylidae and the Coelosomidae. However, a more reasonable classification demands 
further investigations especially on the morphogenetic processes. 

Ruinen (1938) is wrong in transferring Palmarium salinum Gajevskaja, 1925 to the genus 
Trimyema, since Palmarium is illustrated as having an adoral zone of membranelles (Figs 97-101) 
(Borror, 1972). 

Trimyema pur a (Ehrenberg) is listed by Curds (1975) as a species occurring in percolating 
filters and in activated sludge. We suppose that this species has been described as Trichoda pura 
Ehrenberg, 1831, which according to Corliss & Dougherty (1967) is a synonym of Tetrahymena 
pyriformis. 

Lackey (1925) classified Trimyema as female using the latin ending -a for his species T. 
compressa. Since we could not find any greek word comparable to 'myema' from which the name of 
the genus and its sex could be derived we accept Lackey's proposal of the sex. This, however, 
requires the endings of T. claviforme, T. marinum, T. minutum, and T. pleurispirale to be emended 
(see below). 



Key to the species 

la 3 somatic ciliary spirals 2 

Ib Usually more than 3 somatic ciliary spirals 3 

2a Posterior end of body tapered, length 25-65 um T. compressa 

2b Posterior end of body broadly rounded, prominent beak-like pharynx opening, length c. 20 um . 

. T. minuta 

3a Body broadly oval, width c. half length of body 4 

3b Body rather slender, fusiform or oblong, width much less than half length of body ... 5 



THE GENERA ACINERIA, TRIMYEMA AND TROCH1LIOPSIS 209 

4a 4(-6) somatic ciliary spirals restricted to the anterior half of body, length c. 20-45 um . 

T.pleurispiralis 

4b 7 somatic ciliary spirals restricted to the anterior half of body, length c. 25^0 jam, endocommen- 

sally in sea-urchins T. echinometrae 

5a Body club-shaped, thickened in the anterior region and slender in the posterior region, length c. 

40 urn, (not totally reliable species!) T. claviformis 

5b Body not club-shaped . . 6 

6a Shape of body obviously asymmetric, tapered at both ends, anterior pole bent to the right, 

posterior pole bent to the left, peristome measures c. one third of cell length . . T.kahli 

6b Shape of body symmetrical, slender fusiform to slender oblong 7 

7a Body length c. 40 urn, peristome measures c. one fourth of body T. marina 

7b Body length c. 60 um, peristome measures less than one fourth of body (not totally reliable 

species!) .............. T. alfredkahti 



Descriptions of species 
Trimyema compressa Lackey, 1925 

Sciadostoma difficile Kahl, 1926 
Trimyema compressum Kahl, 1933 
Trimyema marinum Faure-Fremiet, 1962 

NEOTYPE-SPECIMENS. Slides (dry silvered and protargol silver impregnated) of neotype-specimens 
have been deposited in the British Museum (Natural History) in London, reference numbers 
1986:5:30:2-3. 

REDESCRIPTION (Figs 59-86, 104-1 13, Table 2). Type species of the genus. Freshwater and marine. 
In vivo about 25-50(-60) x 15-20(-35) urn (Lackey, 1925; Kahl, 1926, 1928, 1931, 1933; Wang & 
Nie, 1935; Liebmann, 1936;Czapik, 1975a; Schmall, 1976; Detcheva 6>f0/., 1981). Body fusiform to 
plump S-shaped, laterally slightly flattened, anterior and posterior end slightly tapered. Dorsally 
and ventrally an inconspicuous ectoplasmatic ridge, the so-called keel (unrecognized by us) (Kahl, 
1926; Wang & Nie, 1935). Macronucleus spherical to slightly oval, located centrally in most 
specimens. Schmall (1976) found it to be more variable, also located posteriorly. Micronucleus 
closely attached to the macronucleus. In protargol impregnated specimens often a second, weakly 
stained macronucleus-like structure, probably a large ingestion vacuole (Fig. 109). Macronucleus 
usually heavily stained, surrounded by dark, slightly curved rods measuring c. 2 um in length and 
0-5 um in width. These aggregated rods look like bacteria. Detcheva et al. (1981), however, 
consider them to be mitochondria, which is not supported by recent studies on other sapropelic 
ciliates (Van Bruggen et al., 1984). Contractile vacuole and its pore located in the region of the last 
ciliary spiral on the right ventro-lateral side. Cytoproct a slit circa 5-10 um long, located in the 
right dorso-lateral surface (Figs 104, 105, 108). Pellicle thin, flexible and deformable, with very 
slight ridges paralleling the longitudinal kinetics. In protargol impregnated specimens these ridges 
appear darkly stained and produce a negative image of the silverline system. Cytoplasm rather 
transparent, contains a lot of refractive long-oval (length about 0-5-1-5 um) granules which are 
also visible in protargol stained specimens. They are most probably the mucocysts described by 
Detcheva et al. (1981). Food vacuoles about 5 um in diameter. Cyclosis pronounced (Lackey, 
1925). Feeds on bacteria but is not dependent on sulphur bacteria (Liebmann, 1947). Moves slowly 
and slightly tremblingly in a straight line or in the arc of a circle rotating on its longitudinal axis 
(Lackey, 1925). Reproduction by transverse fission (Lackey, 1925; Kahl, 1926, 1931; Czapik, 
1975a). 

Somatic cilia 7-9 um, strongly beating, arranged in about 50-60 longitudinal kinetics but more 
commonly viewed as 3 oblique spirals. In the anterior region of these spirals the third, fourth, and 
fifth kinetosomes are paired, constituting the compact field of cilia, consisting of 3 x 4 and 2x2 
cilia, described by Schmall (1976) (Figs 75, 1 13). A short row of about 5-10 cilia on the ventral side 
extends obliquely from the posterior end of the anteriormost somatic spiral to the right. Posterior 




Figs 77-86 Trimyema compressa. 77-79 After Jankowski ( 1 964). 77 From life (inverted). 78 Left lateral 
view of a dry silver impregnated specimen. 79 Lateral view of a dry silvered specimen (inverted). 80-82 
After Czapik (1975a), specimens stained by Chatton's method as modified by Corliss. 80 Scheme 
representing the disposition of the ciliary spirals. 81 Ventro-apical region. 82 Ventral side (Czapik 
called it right side). 83 After Detcheva et al. ( 1 98 1 ), dorsal view (inverted). 84-86 After Faure-Fremiet 
(1962) who identified it erroneously as T. marina. 84 From life. 85 Apical view of Chatton-Lwoff 
impregnated specimen. 86. Infraciliature and silver lines of left dorso-lateral side of a Chatton-Lwoff 
impregnated specimen (Faure-Fremiet interpreted it as left ventro-lateral view). 



THE GENERA ACINERIA, TRIMYEMA AND TROCHILIOPSIS 



211 





95 





212 H. AUGUSTIN, W. FO1SSNER & H. ADAM 

third of body unciliated apart from the caudal cilium measuring about one third to one half of body 
length (Lackey, 1925; Kahl, 1931; Wang & Nie, 1935); it is perhaps involved in the process of 
defecation (Liebmann, 1936) (Figs 62-65). 

Vestibulum circa one third of body length, funnel-shaped. Left half of the oral depression more 
excavated than the right one and, as a consequence, the left margin becomes a thin, transparent 
layer of ectoplasm and forms a cap or hood-like process bordering the vestibulum (Kahl, 1926; 
Wang & Nie, 1935). Cytopharyngeal fibres inconspicuous, rectangular to the entrance of the 
vestibulum. Vestibular kinety 1 a bit longer than vestibular kinety 2. At their anterior ends 4 to 5 
pairs of basal bodies or single basal bodies with parasomal sacs. Vestibular kinety 3 consists of only 
6-7 cilia (Figs 107, 111, 112). In stained specimens somatic as well as vestibular kinetosomes 
appear to be paired (Figs 106-1 1 3) but in fact, the anterior granule is a parasomal sac (Detcheva et 
al., 1981), probably with the exception of the above mentioned compact field. 

About 60 longitudinal silver lines (Czapik, 1975a mentioned 52 lines), connected by transverse 
lines which are located between the somatic ciliary spirals. In front of the anteriormost ciliary spiral 
a circumoral silver line from which a few longitudinal lines extend to the vestibulum forming 
square-like fields at its rim. The longitudinal silver lines fuse at the posterior third forming rough 
meshes (Figs 106, 110). 

OCCURRENCE AND ECOLOGY. First recorded from the sewage disposal of Imhoff tanks in New Jersey 
and later listed as an obligate anaerobe (Lackey, 1925,1938; Noland & Gojdics, 1 967). Very similar 
habitats were reported by Liebmann (1936, 1947, 1951), who found T. compressa regularly in 
waters containing a lot of organic matter and H 2 S, such as in over-loaded percolating filters, in 
Imhoff tanks (S-Sind.ml" 1 and 40 hid. ml" 1 ), in sewers, and at the outfalls of communal waste 
waters. 

Further habitats are the sapropel of ponds near Leningrad (Jankowski, \964a,b), ponds used for 
the treatment of sugar factory wastes (Grabacka, 1973), the plankton of the eutrophic pond 
Toppelsdorfer Weiher' in Bonn (Wilbert, 1969), a small eutrophic lake at Uttendorf/Salzburg 
(Foissner, unpublished), and an arctic tundrapond at Barrow/Alaska (Fenchel, 1975). Detcheva 
( 1 972) and Czapik ( 1 915a,b) listed up Bulgarian and Polish habitats like ponds, lakes, ditches, and 
polluted rivers. Wang & Nie (1935) observed some individuals among decaying organic substances 
taken from Lake Ho Hu. Kahl ( 1 926, 1931,1933) found it in the sapropel, in a cesspool, in sewage, 
and more rarely in the brackish waters of Oldesloe (Kahl, 1928) thus considering it to be of 
freshwater origin. Faure-Fremiet (1962) found it in a rock pool on the French Atlantic coast. 
Tucolesco (\962b) recorded it from the Black Sea and from the saliferous, para-marine Lake 
Tekirghiol in Roumania. According to Sladecek (1972) T. compressa developed in great numbers 
(up to 10,000 ind.ml" *) in a sample of industrial waste water from a textile factory. 



Figs 87-103 Trimyema. 

Figs 87-89 Trimyema marina. 87, 88 After Kahl (1933). 89 After Kahl (1931). 

Figs 90, 91 Trimyema minuta after Kahl (1931), dorsal and left lateral view. 

Fig. 92 Trimyema claviformis after Kahl ( 1 933). 

Figs 93, 94 Trimyema echinometrae after Groliere et al. ( 1 980), protargol silver impregnated specimens 
(inverted), lateral view and ventral view (the latter designated as lateral view, too). 

Figs 95, 96 Trimyema pleurispiralis after Borror (1972). 95 Left ventro-lateral view. 96 Anterior pole 
with anteriormost somatic ciliary spiral (at the left) and with vestibular ciliature. 

Figs 97-101 Palmarium salinum Gajevskaja after Ruinen (1938). 97, 98 Normal form, ventral and 
dorsal view. 99 Slender (abnormal?) form. 100, 101 Peristome, ventral and lateral view. 

Fig. 102 Trimyema alfredkahli after Tucolesco ( 1 962a), left lateral view. 
Fig. 103 Trimyema kahli after Tucolesco ( 1 962o), left lateral view. 



THE GENERA ACINERIA, TRIMYEMA AND TROCHILIOPSIS 



213 




107 



Figs 104-109 Trimyema compressa, originals, scale = 20 um each. 104 Left ventro-lateral view, from 
life. 105 Right dorso-lateral view of an S-shaped individual. CP, cytoproct. 106 Dorsal view of a dry 
silvered specimen. 107 Vestibular ciliature revealed by Fernandez-Galiano's method. The shape of the 
vestibular kinetics has been slightly deformed by preparation; they are less curved in life. Vl-3, 
vestibular kinetics 1-3. 108, 109 Ventral and dorsal view of a protargol silver impregnated specimen 
amended with details from individuals impregnated with Fernandez-Galiano's method. CVP, 
contractile vacuole pore. 



214 



H. AUGUSTIN, W. FOISSNER & H. ADAM 




^^Y** l **^s. ' V 

X. * *i* ^T^k- ' 







110 



111 




* -^ * . * * 

. 



112 



113 



Figs 110-113 Trimyema compressa. 110 Silverline system revealed by the dry silver impregnation 
technique, dorsal view. 111-113 Specimens stained by Fernandez-Galiano's impregnation technique. 
Ill Left side with the three ciliary spirals and apex with vestibular ciliature consisting of two long rows 
and one short row of cilia (arrow). 112 Dorsal view, arrow indicates the three vestibular kinetics. 113 
Ventral view, arrow indicates the isolated basal bodies at the anteriormost region of the vestibular 
kinetics. 



THE GENERA ACINERIA, TRIMYEMA AND TROCHILIOPSIS 2 1 5 

Bick (1968, 1972) gave the most detailed ecological characterization. T. compressa is an 
outstanding indicator of polysaprobity (Liebmann, 1951), isosaprobity and even metasaprobity 
(Sladecek, 1973) and occurs in waters receiving fresh manure and sewage, or waste waters contain- 
ing cellulose material (paper mill outlets, etc.). The species seems to prefer conditions with low 
ammonia content, i.e. conditions prevailing during the decay of cellulose and other material poor 
in nitrogenous compounds. The saprobiological evaluation is indicated by Sladecek (1972): x = 0, 
o = 0, = 0, a = 0, p= 10, G = 5, s = 5-3 (E, H 2 S). 

REMARKS. This species differs from T. minuta particularly by the tapered posterior end. It can easily 
be distinguished from the other species by its having only three somatic ciliary spirals. Faure- 
Fremiet (1962) observed an abundant population of Trimyema (Figs 84-86) and identified it as T. 
marina although it was of an ovoid and stocky form which was not described by Kahl (1931, 1933, 
1935). On the contrary this author later stated that T. marina is usually one third to one half more 
slender than he drew it in 1 93 1 (Kahl, 1931, Fig. 89; Kahl, 1933, Figs 87,88). Thus we suppose that 
Faure-Fremiet worked on T. compressa. 

Table 2 Biometrical characterization of Trimyema compressa 

Character 1 x M SD SE CV Min Max n 



Body, length 


39-05 


39 


5 


4-58 


1-03 


11-7 


32-0 


47-0 


20 


Body, width 


22' 


30 


23' 





3-01 


0-67 


13-5 


17-0 


26-0 


20 


Macronucleus, length 


11 


05 


11 





1-57 


0-35 


14-2 


9 





14-0 


20 


Macronucleus, width 


9 


35 


9 


'5 


1-50 


0-34 


16-0 


6-0 


12-0 


20 


Number of vestibular ciliary rows 


3' 


00 


3 





0-00 


0-00 


0-0 


3 





3-0 


20 


Number of somatic ciliary rows 


3 


00 


3 





0-00 


0-00 


0-0 


3 





3-0 


20 


Number of caudal cilia 


1 


00 


1 





0-00 


0-00 


0-0 


1 





1-0 


20 


Distance from apex to posterior end of vestibulum 


9 


80 


10-0 


1-88 


0-42 


19-2 


7 





15-0 


20 


Distance between posterior end of body and posterior 
























end of ciliary spirals 


11 


00 


11 





1-78 


0-40 


16-2 


7 





15-0 


20 



1 See footnote Table 1 



Trimyema a#ra/M Tucolesco, \962a 

DIAGNOSIS (Fig. 102). Marine. About 60 jim. Body oblong and slender, slightly tapering anteriorly 
and posteriorly. Oral apparatus particularly small, bounded at the right margin by a conspicuous 
dilatation. Macronucleus spherical. Cilia long and fine. According to Tucolesco's figure ciliary 
spirals cover nearly the whole body, which contrasts his description. Caudal cilium longer than half 
body length. 

OCCURRENCE AND ECOLOGY. Found in an abundant population in a mixed polysaprobic culture 
taken from the Black Sea in March 1955. 

REMARKS. This species has remained unmentioned since description. It can perhaps be dis- 
tinguished from T. marina by its oblique orientation of the oral apparatus, which is stressed by 
Tucolesco (1962#), and by its larger size. However, synonymy cannot be excluded. 

Trimyema claviformis Kahl, 1933 
Trimyema claviforme Kahl, 1933 

DIAGNOSIS (Fig. 92). Marine. Circa 40 urn. Body club-shaped. Posterior third of body unciliated. 
OCCURRENCE AND ECOLOGY. Found in sapropelic habitats of Sylt and Kiel (Germany). 



216 H. AUGUSTIN, W. FOISSNER & H. ADAM 

REMARKS. Very insufficiently described. With exception of the unciliated tapering posterior third of 
body identical with T. marina. Even Kahl (1935) noted that he established this species with some 
doubt. Thus, synonymy cannot be excluded. 

Trimyema echinometrae Groliere, Puy torac & Grain, 1 980 

DIAGNOSIS (Figs 93, 94). Marine. Living endocommensally in sea-urchins. About 31 (27-40) x 17 
(1 3-20) urn. Body peg-top like. Macronucleus spherical, 5-7-5 urn in diameter, posteriorly located. 
Micronucleus not visible. 60 to 70 longitudinal somatic kinetics. Cilia distributed in 7 parallel 
spirals in the anterior half of body. Three vestibular kinetics very similarly arranged as in T. 
compressa. 

OCCURRENCE AND ECOLOGY. Found in the sea-urchins Diadema antillarum and Echinometra 
lucunter from the Gulf of Mexico and the Gulf of Guadeloupe. Housing together with other 
commensal species like Biggaria echinometris, Metanophrys elongata and Metopus circumlabens 
(Groliere et al., 1980). Perhaps already Profant (1966) observed this species, since he mentioned 
Trimyema sp. to be a ciliate inhabiting echinoids in the Eastern Pacific Ocean. 

REMARKS. T. echinometrae is a reliable species. It differs from the other members of the genus in the 
number of ciliary spirals. The figures, however, are obviously inverted, because in the genus 
Trimyema the spirals run the other way round. Furthermore, the identification is impeded by the 
missing drawing from life. 

Trimyema kahli Tucolesco, \962a 

DIAGNOSIS (Fig. 103). Para-marine. About 36-40 urn. Body conspicuously asymmetric, inverted 
S-shaped. Peristome in the anterior third of body. Macronucleus spherical, usually located in the 
middle of the cell. Contractile vacuole close behind the middle of body. Cilia long and fine. Ciliary 
spirals extending to the posterior pole. Caudal cilium almost rigid, bent to the left. 

OCCURRENCE AND ECOLOGY. Polysaprobic, found constantly in the para-marine Roumanian Lake 
Tekirghiol (Tucolesco, \962a,b). 

REMARKS. This species has remained unmentioned since 1962. However, from its general 
appearance it seems to be a reliable but insufficiently described species. 

Trimyema marina (Kahl, 1931) 

Sciadostoma marinum Kahl, 1931 
Trimyema marinum Kahl, 1933 

DIAGNOSIS (Figs 87-89). Marine. About 40 urn. Slender fusiform to slender oblong (4 : 1). In the 
original figure (Fig. 89) similar to T. compressa but later figured and redescribed with 5-6 ciliary 
spirals (Figs 87, 88). 

OCCURRENCE AND ECOLOGY. Repeatedly observed in putrid water of the North and East Sea (Sylt, 
Kiel) and in salt-water from Oldesloe (Kahl, 1931, 1933, 1935). 

REMARKS. Kahl (1931) considered T. marina to be a separable species because he never found 
similar forms among numerous populations of the freshwater form of T. compressa. Later he 
thought that two forms of this species probably exist and erected the species T. claviformis (Kahl, 
1933) which, however, is not a totally reliable species (Kahl, 1935). We consider this species and 
T. alfredkahli perhaps to be junior synonyms of T. marina. 

Trimyema minuta nov. comb. 
Sciadostoma minutum Kahl, 1931 

DIAGNOSIS (Figs 90, 91). Freshwater and marine. About 20 um. Rounded posterior and a 
prominent beak-like pharynx-opening. Ectoplasmatic ridge (keel) more pronounced than in T. 



THE GENERA ACINERIA, TRIMYEMA AND TROCHILIOPSIS 217 

compressa, extending from the beak-like pharynx-opening over the back to the posterior. Cilia 
longer and more rigid than in T. compressa. 

OCCURRENCE AND ECOLOGY. This species was found together with T. compressa and was first 
considered as a modification, but once an abundant population occurred in a ditch contaminated 
with liquid manure (Kahl, 1931). Wenzel (1961) observed T. minuta in the sponge Halichondria 
panicea from the Gulf of Naples. Tucolesco (19626) recorded it twice from old, mixed infusions of 
the para-marine Roumanian Lake Tekirghiol. 

REMARKS. Kahl (1931) doubted the species status of this form and did not mention it again in his 
publication of the year 1935. Further investigations are necessary. 

Trimyema pleurispiralis Borror, 1972 

DIAGNOSIS (Figs 95, 96). Marine. About 20-44 x 16-23 (usually less than 20) urn. Shape of 
prepared individuals egg-like, circular in cross section (Fig. 95). Macronucleus spherical, central. 
Micronucleus not observed. Cytoproct an elongated (approximately 8 urn) slit near posterior pole, 
lying in the same latitude as cytostome and suture at ends of ciliary spirals. Contractile vacuole 
pore not observed. Except for elongated caudal cilium, all somatic cilia restricted to anterior half 
of cell, arranged in at least four spirals (a few individuals possess a partial or even complete 
fifth spiral, and even a few cilia of a sixth spiral). Outer vestibular kinety in a semicircle 
dipping posteriorly into vestibulum and terminating near cytostome. Inner vestibular kinety with 
three regions: (1) anteriormost two isolated tufts of approximately five cilia each, (2) a row of 
kinetosomes closely paralleling the outer kinety, extending from the tufts down to cytostome, (3) 
posteriormost a J-shaped field of cilia. As already mentioned, this interpretation of the oral 
structure is a little erroneous and incomplete. 

OCCURRENCE AND ECOLOGY. Like the other species of this genus T. pleurispiralis is bacterivorous 
and occurred only irregularly in New Hampshire tidal salt marshes (Borror, 1972). 

REMARKS. This species differs from the other members of the genus in number and location of 
ciliary spirals, which are restricted to the anterior half of body. Unfortunately, Borror (1972) did 
not give a drawing from life. Thus, the real body shape is unknown. Redescription is needed. 



Genus TROCHILIOPSIS Penard, 1922 

DIAGNOSIS. Microthoracidae Wrzesniowski, 1870 with cytostome in the anterior third of body. 
Three preoral kinetics subapically on the left body side. Somatic kinetics from either side terminate 
near the pointed beak-like region formed by the oral structures. Apex smooth. Rightmost somatic 
kinety of the right side interrupted. Contractile vacuole located almost centrally. Freshwater, 
polysaprobic. 

TYPE-SPECIES. TrochiliopsisopacaPenard, 1922. 

REMARKS. Trochiliopsis shows many characters which are very likely homologous to genera of the 
family Microthoracidae Wrzesniowski 1870 according to the classification of Foissner (19856). 
Thus, a separation of Trochiliopsis at the familial level as suggested by Jankowski (1975) is not 
justified (Compare Corliss, 1979; Curds, 1982). On the contrary, the organization of Trochiliopsis, 
especially the general appearance of the infraciliature and the location and structure of the oral 
apparatus, allows a classification close to the genus Stammeridium. These similarities might have 
induced Kahl (1931) to synonymize Trochiliopsis with Trichopelma Levander and Leptopharynx 
Mermod. There are just sufficient differences in the location of the preoral kinetics, the paroral 
membrane, and the shape of the anteriormost region for separating these two genera. Further- 
more, by a trivial twist of some organelles of Trochiliopsis, the typical organization of the genus 
Stammeridium can be achieved (Figs 126, 127): The preoral kinetics move to the apex between 
serrated processes, the paroral membrane gets located obliquely to the longitudinal axis and the 
contractile vacuole moves close to the ventral side. 



218 H. AUGUSTIN, W. FOISSNER & H. ADAM 

Key to the genera of Microthoracina Jankowski 1967 (based on Foissner 1985&) 

la Microthoracina with somatic cirri-like organelles, fusiform extrusomes, and wide-meshed silver- 
line system .......... (Discotrichidae) Discotricha 

1 b Microthoracina with normal cilia, anchor-like extrusomes, and granular or fine-meshed silverline 
system ................. 

2a Microthoracina with more than 10 uninterrupted somatic kinetics 

(Pseudomicrothoracidae) Pseudomicrothorax 

2b Microthoracina with fewer than 1 0, usually 6 partly interrupted somatic and three preoral kinetics 

(Microthoracidae) 

3a Oral apparatus ventrally in the posterior third of body Microthorax 

3b Other .... 

4a Oral apparatus ventrally between middle and posterior third of body, body more or less oblong . 

Drepanomonas 

4b Other . . 

5a Oral apparatus between middle and anterior third of body, rightmost somatic kinety or right side 
uninterrupted, preoral kinetics run in distinct furrows from the ventral to the right body side and 

form a keel Leptopharynx 

5b Other . ... 

6a Rightmost somatic kinety of right side interrupted, preoral kinetics run anterior-posteriorly on the 
left side of the body, paroral membrane circa half body length .... Trochlliopsis 

single species: Trochiliopsis opaca 
6b Preoral kinetics apically in furrows, apex distinctly serrated, paroral membrane shorter than a 

third of body running obliquely to the longitudinal axis Stammeridium 

single species: Stammeridium kahli 



Description of species 
Trochillopsis opaca Penard, 1 922 

Trichopelma opaca Kahl, 1931 
Leptopharynx opaca Detcheva, 1972 

NEOTYPE-SPECIMENS. Slides (protargol silver impregnated and dry silvered) of neotype-specimens 
have been deposited in the British Museum (Natural History) in London, reference numbers 
1986:5:30:4-5. 

REDESCRIPTION (Figs 114-131, Table 3). Type species of the genus. Freshwater. In vivo circa 
30-40(-50) x 17-20 um. Body outline oval, anteriorly curved slightly to the ventral side terminat- 
ing in a pointed beak-like region (peak). Body strongly compressed laterally (circa 2:1). Somatic 
kinetics in deep, crenelated furrows, which terminate near the oral peak. Macronucleus spherical, 
more or less centrally located, in vivo hardly discernible. Micronucleus closely attached to the 
macronucleus. Contractile vacuole centrally located, close to the right lateral surface, diameter 
about 4 um; contractile vacuole pore at the end of the paroral membrane. Cytoproct slightly 
posterior to the contractile vacuole pore, visible as black line in dry silvered specimen (Fig. 130). 
Pellicle rigid, colourless, opaque. Extrusomes about 3 um, fusiform, scattered over the whole body 
in the ribs between the furrows, show four anchor-like processes at the distal end in the exploded 
phase. Probably feeds on bacteria, but no food vacuoles were found. Slow, trembling and swaying 
movements. 

Length of cilia 8-10 um. Six somatic kinetics (Kl-6), three preoral kinetics (Pl-3), and a short 
x-kinety (Figs 124, 125). Kl anterior with 8-10, posterior with 4, K2 anterior with 2, posterior with 
5-6, K3 anterior with 12-16, posterior with 5-8, K4 (anterior) with 6-8, K5 anterior with 3-4, 
posterior with 2, K6 with 10-12 kinetosomes. At the end of K4 and in the middle of K5 sometimes a 
single unciliated kinetosome, respectively. Basal bodies of Kl-5 mostly paired, K6 always with 
single kinetosomes. Preoral kinety 1 with 4-5 pairs, preoral kinety 2 constantly with 5 singles, and 
preoral kinety 3 constantly with 7 singles, x-kinety with 1-2 paired basal bodies located left of the 
posterior end of the paroral membrane (Figs 120-126). 



THE GENERA ACINERIA, TRIMYEMA AND TROCHILIOPSIS 



219 



116 




114 



115 



117 



118 



119 




120 



121 



122 



123 



K1 




K3 



125 





126 



127 



Figs 114-126 Trochiliopsisopaca, scale = 10 urn each. 114-117 After Penard( 1922). 114, 115 Right and 
left lateral view. 116 View from the apex. 117 Extrusomes with 2, 3, and 4 processes. 118, 119 After 
Kahl (1931), right and left side. 120-123 Originals, from life and protargol silver stained specimens, 
right and left side respectively. CP, cytoproct. 124, 125 Schematized organization of T. opaca, right 
and left lateral view. Kl-6, somatic kinetics 1-6; M, adoral membranelles; PM, paroral membrane; 
Pl-3, preoral kinetics 1-3; x-K, x-kinety 126 Probable evolution of Stammer idium from Trochiliopsis. 

Fig. 127 Schematic organization of the genus Stammeridium (after Foissner, 1985). 



220 



H. AUGUSTIN, W. FOISSNER & H. ADAM 




/' ' i 




128 




129 




130 




131 



Figs 128-131 Trochiliopsis opaca. 128, 129 Protargol silver impregnated specimens, right and left side. 
130, 131 Dry silver impregnated specimens, right and left side. 



Probably only two adoral membranelles, located at the oral peak. Anterior adoral membranelle 
most likely built up by two rows, posterior one probably by three rows of kinetosomes. Paroral 
membrane with 8-9 paired basal bodies (Figs 120, 122, 124). Cyrtos invisible in life even with 
interference contrast, but slightly impregnated with protargol silver. 

Silverline system granular or very fine-meshed. 

OCCURRENCE AND ECOLOGY. Penard (1922) found few individuals between dead leaves of the 'swan 
pond' at Ariana ('. . . a 1'etang des Cygnes', Switzerland). Kahl (1931) noted T. opaca sporadically 
in the sapropel and sometimes numerously in sapropelic infusions of Glyceria. Lackey (1938) 
recorded it once from a polluted stream, twice from a trickling filter, and five times from an 
activated-sludge chamber. Noland & Gojdics (1967) mentioned that T. opaca occurs when the 
sludge has reached the finely particulate stage and the bacteria in it are well distributed. Detcheva 



THE GENERA ACINERIA, TRIMYEMA AND TROCHILIOPSIS 



221 



(1972) listed some Bulgarian habitats, namely a pond in the surroundings of the village Bosnek in 
the Witoscha mountains, a marshy meadow in the vicinity of the village Kasitschene near Sofia, 
and a river in the Wrabniza quarter of Sofia. Apart from in activated sludge, we found this species 
once in the polysaprobic zone of a heavily polluted river (Ager near Lenzing, Upper Austria). 
These localities suggest T. opaca to be a good indicator of heavily polluted (polysaprobic) 
conditions. It might also have some tolerance of H 2 S. 



Table 3 Biometrical characterization of Trochiliopsis opaca 



Character 1 



M SD SE CV Min Max n 



Body, length 


25' 


66 


26-0 


1-12 


0-37 


4-4 


24-0 


27-0 


9 


Body, width 


13 


'22 


13-0 


0-83 


0-28 


6-3 


12-0 


15-0 


9 


Macronucleus, length 


6 


33 


6-5 


0-35 


0-12 


5-6 


6-0 


7-0 


9 


Macronucleus, width 


6 


11 


6-0 


0-42 


0-14 


6-8 


5-5 


7-0 


9 


Distance from apex to the beginning of macronucleus 


12 


22 


12-0 


1-30 


0-44 


10-6 


10-0 


14-0 


9 


Micronucleus, length 


1 


62 


1-6 


0-30 


0-10 


18-7 


1-2 


2-0 


9 


Micronucleus, width 


1 


51 


1-5 


0-31 


0-10 


20-8 


1-0 


1-8 


9 


Number of kinetosomes of paroral membrane 


17 


78 


18-0 


0-67 


0-22 


3-7 


16-0 


18-0 


9 


Number of kinetosomes of anterior kinety 1 


8 


22 


8-0 


0-67 


0-22 


8-1 


8-0 


10-0 


9 


Number of kinetosomes of posterior kinety 1 


4-00 


4-0 


0-00 


0-00 


0-0 


4-0 


4-0 


9 


Number of kinetosomes of anterior kinety 2 


2' 


00 


2-0 


0-00 


0-00 


0-0 


2-0 


2-0 


9 


Number of kinetosomes of posterior kinety 2 


5 


89 


6-0 


0-33 


0-11 


5-7 


5-0 


6-0 


9 


Number of kinetosomes of anterior kinety 3 


13 


11 


12-0 


1-45 


0-48 


11-1 


12-0 


16-0 


9 


Number of kinetosomes of posterior kinety 3 


6 


11 


6-0 


0-93 


0-31 


15-2 


5-0 


8-0 


9 


Number of kinetosomes of kinety 4 


6 


67 


6-0 


0-87 


0-29 


13-0 


6-0 


8-0 


9 


Number of kinetosomes of anterior kinety 5 


3 


11 


3-0 


0-33 


0-11 


10-7 


3-0 


4-0 


9 


Number of kinetosomes of posterior kinety 5 


2 


00 


2-0 


0-00 


0-00 


0-0 


2-0 


2-0 


9 


Number of kinetosomes of kinety 6 


10 


33 


10-0 


0-71 


0-24 


6-8 


10-0 


12-0 


9 


Number of kinetosomes of the x-kinety 


3 


78 


4-0 


0-67 


0-22 


17-6 


2-0 


4-0 


9 


Number of kinetosomes of preoral kinety 1 


9 


89 


10-0 


0-33 


0-11 


3-4 


9-0 


10-0 


9 


Number of kinetosomes of preoral kinety 2 


5 


00 


5-0 


0-00 


0-00 


0-0 


5-0 


5-0 


9 


Number of kinetosomes of preoral kinety 3 


7 


00 


7-0 


0-00 


0-00 


0-0 


7-0 


7-0 


9 



'See footnote Table 1 



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Manuscript accepted for publication 26 May 1986 






British Museum (Natural History) 

The birds of Mount Nimha, Liberia 

Peter R. Colston & Kai Curry-Lindahl 



For evolution and speciation of animals Mount Nimba in Liberia, Guinea and the Ivory Coast is 
a key area in Africa representing for biologists what the Abu Simbel site in Egypt signified for 
archaeologists. No less than about 200 species of animals are endemic to Mount Nimba. Yet, this 
mountain massif, entirely located within the rain-forest biome, is rapidly being destroyed by 
human exploitation. 

This book is the first major work on the birds of Mount Nimba and surrounding lowland 
rain-forests. During 20 years (1962-1982) of research at the Nimba Research Laboratory in 
Grassfield (Liberia), located at the foot of Mount Nimba, scientists from three continents have 
studied the birds. In this way Mount Nimba has become the ornithologically most thoroughly 
explored lowland rain-forest area of Africa. 

The book offers a comprehensive synthesis of information on the avifauna of Mount Nimba 
and its ecological setting. During the 20 years period of biological investigations at Nimba this in 
1962 intact area was gradually opened up by man with far-reaching environmental consequences 
for the rain-forest habitats and profound effects on the birds. Therefore, the book provides not 
only a source of reference material on the systematics, physiology, ecology and biology of the 
birds of Mount Nimba and the African rain-forest, but also data on biogeography in the African 
context as well as conservation problems. Also behaviour and migration are discussed. At 
Nimba a number of migrants from Europe and/or Asia meet Afrotropical migratory and 
sedentary birds. 

Professor Kai Curry-Lindahl has served as Chairman of the Nimba Research Laboratory and 
Committee since its inception in 1962. Peter Colston is from the Subdepartment of Ornithology, 
British Museum (Natural History), Tring, and Malcolm Coe is from the Animal Ecology 
Research Group, Department of Zoology, Oxford. 

1986, 129pp. Hardback. 565 00982 6 17.50. 



Titles to be published in Volume 52 



Miscellanea 

A revision of the Suctoria (Ciliophora, Kinetofragminophora) 5. The Paracineta 
and Corynophora problem. By Colin R. Curds 

Notes on spiders of the family Salticidae 1. The genera Spartaeus, Mintonia and 
Taraxella. By F. R. Wanless 

Mites of the genus Holoparasitus Oudemans, 1936 (Mesostigmata: Parasitidae) in 
the British Isles. By K. H. Hyatt 

The phylogenetic position of the Yugoslavian cyprinid fish genus Aulopyge Heckel, 
1841, with an appraisal of the genus Barbus Cuvier & Cloquet, 1816 and the 
subfamily Cyprininae. By Gordon J. Howes 

Revision of the genera Acineria, Trimyema, and Trochiliopsis (Protozoa, 
Ciliophora). By H. Augustin, W. Foissner & H. Adam 

The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) with a 
systematic review, a synopsis of Pipistrellus and Eptesicus, and the descriptions of 
a new genus and subgenus. By J. E. Hill & D. L. Harrison 

Notes on some species of the genus Amathia (Bryozoa, Ctenostomata). 

By P. J. Chimonides 



Printed in Great Britain by Henry Ling Ltd., at the Dorset Press. Dorchester. Dorset 



Bulletin of the 

British Museum (Natural History) 



The baculum in the Vespertilioninae 
(Chiroptera: Vespertilionidae) with a 
systematic review, a synopsis of Pipistrellus 
and Eptesicus, and the descriptions of a nev* 
genus and subgenus 



J. E. Hill & D. L. Harrison 



Zoology series Vol52 No 7 30 July 1987 



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Assistant Editor : Mr C. G. Ogden 



ISBN 0565 05031 1 

I SSN 0007- 1 498 Zoology series 

Vol 52 No. 7 pp 225-305 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD Issued 30 July 1 987 



(NATURAL HISTORY) 

,-;c? 
The baculum in the Vespertilioninae (Chiroptera: 

Vespertilionidae) with a systematic review, a synopsis 
of Pipistrellus and Eptesicus, and the descriptions of a 
new genus and subgenus 

J. E. Hill 

Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD 

D. L. Harrison 

Harrison Zoological Museum, Bowerwood House, St Botolph's Road, Sevenoaks, Kent 
TN133AL 



Contents 

Synopsis 

Introduction 

Functional and systematic significance of the baculum 

Nature and scope of this study 

Materials and methods 

Authorship and responsibility 

The baculum of Pipistrellus 

The baculum of Eptesicus 

The baculum in other Vespertilioninae 

Systematic considerations ..... 

Genus Pipistrellus Kaup, 1829 

Genus Eptesicus Rafi nesque, 1 820 

The status of the 'Nycticeini'. 

The classification of the Vespertilioninae 
Zoogeographical considerations 

Conclusions 

Addendum 

References 

Appendix 1 . Specimens examined 

Tables 1-3 

Figures 1-22 




Synopsis 

Current classification of the Vespertilioninae rests chiefly on a suite of mainly adaptive characters, among 
which facial shortening throughout the subfamily with consequent changes in the structure, size, relative 
position and number of the incisive and premolar teeth features prominently. Such characters may not 
necessarily reflect relationships or phyletic diversity, and sometimes do not serve properly to distinguish the 
genera that they purport to define, as in the distinction of Pipistrellus and Eptesicus, where generic boundaries 
remain unclear. The search for possibly less strongly adaptive features suggested the possibility that the 
morphology of the os penis or baculum might prove valuable in the study of the systematics of these genera 
and perhaps in the subfamily as a whole. 

This paper reviews earlier studies of the baculum in the Chiroptera and their relevance to systematics in the 
Order, with an examination of its gross morphology throughout the Vespertilioninae, especial attention being 
given to species currently allocated either to Pipistrellus or to Eptesicus. A synoptic review of the species 



Bull. Br. Mus. nat. Hist. (Zool.) 52 (7): 225-305 



Issued 30 July 1987 



225 



226 J. E. HILL & D. L. HARRISON 

content of these genera is presented, with the recognition and definition of subgenera and included species 
groups: three such (pumilus, capensis and tenuipinnis) currently referred to Eptesicus on dental grounds seem 
instead more closely related to Pipistrellus to which they are here transferred. One subgenus of Pipistrellus is 
described as new (p. 250). 

The Vespertilioninae as a whole display a wide range of bacular variation, which falls into two major and 
several minor groups. This has suggested a revision of the current classification of the subfamily, combining 
bacular features with those conventionally in use. Bacular morphology provides a clear indication that 
the 'Nycticeini' (or 'Nycticeiini') is an artificial grouping and that the genus Nycticeius as presently under- 
stood is composite. Currently it is held to include two species, the North American humeralis and the African 
schlieffenii: these are here thought to be sufficiently characterised to justify generic separation and a new 
generic name is proposed for schlieffenii (p. 254). 

A suggested classification of the subfamily is presented, with a tabulated review of earlier classifications; 
possible relationships between the constituent genera are discussed and the zoogeography of the bacular types 
within the subfamily is examined. 



Introduction 

A penial bone is known to occur among mammalian Orders in the Insectivora, Chiroptera, 
Primates, Rodentia and Carnivora. Variously called the os penis, os priapi or os glandis, it was first 
named the baculum by Thomas (191 5a), the corresponding structure in the female, the os clitoridis, 
being later called the baubellum by Shortridge (1934: 327, footnote). The features of the baculum 
have been used extensively in attempts to determine phyletic relationships at various systematic 
levels (Patterson & Thaeler, 1982). Thomas (loc. cit.), for example, suggested that the baculum 
might provide evidence valuable in the subfamilial classification of the Sciuridae and indeed 
pointed out that in this connection there were no bacular features to support the association of the 
dwarf squirrels in a separate subfamily, the Nannosciurinae. More commonly, bacular features 
have been used to indicate or determine relationships within genera in the Sciuridae, among New 
World rodents, and in the Mustelidae. Such characteristics have been employed in species descrip- 
tions, especially where bacular variation is pronounced, and also for age determination. Numerous 
examples of these uses of the baculum are summarised by Patterson & Thaeler (loc. cit.) while Burt 
(1960) gave an account of the earlier of such studies. Similar early accounts of the baculum in the 
Chiroptera are reviewed by Hamilton (1949). 

The presence of a baculum in some at least of the Chiroptera has been long established. 
Daubenton (1760) described and illustrated (in part) the baculum of Nyctalus noctula and 
Blainville (1840) similarly studied the baculum of Rhinolophus fermmequinum, R. hipposideros, 
Vespertilio murinus and again of Nyctalus noctula, the latter author providing perhaps the first 
accurate and quite detailed drawings of this structure. Later workers such as Ercolani (1868), 
Robin (1881), Gilbert ( 1 892), Rauther ( 1 903), Gerhardt ( 1 905) and Chaine ( 1 926) provided further 
details of penial and bacular morphology in the Chiroptera, Chaine in particular discussing and 
illustrating the baculum in several species and to some extent summarising earlier work in the field. 
However, none attempted to use the structure of the baculum for systematic purposes. 

The first use of the baculum in chiropteran systematics appears to be by Thomas (\9\5b) who 
employed bacular characteristics in defining the species of Nyctophilus. This worker clearly fore- 
saw the value of bacular features in the definition of some at least of the species of bats, beginning 
from that time a collection of vespertilionid bacula at the British Museum (Natural History) 
although subsequently making little use of the material that was accumulating, except in 1928 
employing bacular characters to separate Indo-Chinese species of Pipistrellus (Thomas, 19280, K). 
Since then the baculum has been utilised in a variety of taxonomic studies of bats, for example by 
Krutzsch (1959, 1962) and Lanza (1969) to examine its value in indicating relationships in the 
Megachiroptera, by Topal (19700) in determining the affinities of la, or by Heller & Volleth (1984) 
as an indicator of relationship among the species of Pipistrellus and Eptesicus. The baculum of 
Plecotus was found valuable by Lanza (1960) in discriminating between P. auritus and P. wardi 
( = P. austriacus): the subsequent use of the baculum in distinguishing these species is summarised 
by Corbet (1964). Genoways & Jones (1969) found that bacular features distinguished closely 



VESPERTILIONINE SYSTEMATICS 227 

related species of North American Myotis, LaVal (1973#) employing bacular characters for the 
same purpose among the Neotropical species of this genus. 

The emphasis placed on bacular characters in chiropteran systematics is perhaps best illustrated 
by the number of studies devoted chiefly to bacular structure, often on a regional or faunal basis, as 
for instance the work by Hamilton (1949) and Krutzsch & Vaughan (1955) on North American 
species, by Brown et al. (1971) on Neotropical bats, by Topal (1958) on central European species, 
by Bhatnagar (1967), Agrawal & Sinha (1973), Sinha (1976) and Khajuria (1979, 1980, 1982) on 
Indian bats, or by Wassif & Madkour (1972) and Wassif, Madkour & Soliman (1984) on Egyptian 
bats. Bacula are sometimes studied in discrete taxonomic groupings, as for example those of New 
World molossids by Brown (1967) or of Malaysian Hipposideros by Zubaid & Davison (in press). 
Thus among the Chiroptera the baculum has been employed as a source of taxonomic features at 
several systematic levels, but primarily to indicate degrees of relationship or for separation at the 
specific and sometimes the generic grades, or especially for distinguishing closely related, often 
sympatric species whose conventional morphological characters are otherwise very similar, as in 
Myotis and Plecotus. 



Functional and systematic significance of the baculum 

Conflicting hypotheses for bacular variation were reviewed in detail by Patterson & Thaeler (1982). 
These authors proposed that among rodents at least the probability was that the baculum has a 
precise reproductive purpose and functions primarily as a device contributing to species isolation. 
Bacular differences among closely related taxa might well then take an exaggerated form. As such, 
the baculum would be therefore a poor basis for supra-specific classification, but an excellent 
structure for species diagnosis. Thus they would not consider a phyletic basis for bacular variation 
to be appropriate. They admit, however, that while in some rodent groups there are patterns of 
bacular morphology that do not agree with phyletic divergence as indicated by other morpho- 
logical features, there exist also gross patterns of bacular variation in other groups that do in fact 
conform with accepted phyletic relationships. Indeed, they remarked that there can be little doubt 
that the baculum exhibits phyletic weight and consequently may serve as a valuable taxonomic 
tool. Moreover, taxa that differ in external and cranial characters may have similar bacula, while 
others that are similar in such features may exhibit highly distinctive genitalia. Patterson & 
Thaeler (loc. cit.) suggested that although bacular morphology reflects phyletic history on a gross 
scale, discordance between patterns of bacular and phyletic divergence supports a functional 
interpretation of bacular variation, especially at the species level. 

Similarly, opinions vary as to the value of bacular morphology in chiropteran systematics. 
Hamilton ( 1 949) examined the baculum in North American vespertilionids and concluded that in 
this family the baculum was useful in defining relationships when considered with skull and other 
skeletal characteristics. Thus he was able to suggest that the close similarity between the bacula of 
Myotis (Fig. 19i, j) and Pizonyx (Fig. 19k) indicated their close relationship, and that the dissimi- 
larity between the bacula of Pipistrellus subflavus (Fig. 2d) and P. Hesperus (Fig. 8d) suggested 
generic or at least subgeneric difference. This author also noted that in most instances among 
North American vespertilionids there were marked generic differences in the baculum. He con- 
sidered that further study was needed to determine the usefulness of the baculum in chiropteran 
systematics and that with time and sufficient material the bone might be utilised in classification. 
These conclusions were reinforced by Krutzsch & Vaughan (1955) who examined the bacula of 
further North American species. They remarked that in the case of those that are closely related 
the baculum can serve as a criterion in judging relationship when other clear cut distinguishing 
characters are lacking. These authors found bacular variation in closely related bat species to be 
chiefly in shape, detail of outline, and gross size: their study led to the belief that in at least some 
superficially similar species well marked and consistent bacular differences reinforced the more 
subtle external and cranial dissimilarities. 

Krutzsch (1959) accepted the view that the baculum can provide additional evidence for classifi- 
cation, or, in the absence of other clearly defined characters, can serve as a criterion in judging 



228 J. E. HILL & D. L. HARRISON 

relationship. He added in relation to the Pteropodidae that by virtue of its relative simplicity and 
structural stability the baculum might well serve to help place entities of doubtful relationship in 
their natural position, although they might be otherwise morphologically contradictory. Among 
pteropodids he found that infrageneric differences in the baculum involved minor details of shape, 
outline and size. Genera, however, might be separated by more profound differences. His study 
suggested that although within the genus well marked and consistent differences existed between 
the bacula of individual species, there was nevertheless a basic similarity in pattern throughout the 
genus, leading to the suggestion that marked variants from this morphological standard in a single 
genus might provide grounds for a reappraisal of the affinities of the variant. A further study 
(Krutzsch, 1962) confirmed these opinions, especially in the broad agreement of bacular variation 
in pteropodids with the taxonomic arrangement of this family by Andersen (1912) and by Tate 
(1942ft). Krutzsch concluded that strong intrageneric similarities exist among the bacula of ptero- 
podids, but that representative bacula of different genera differ distinctly: although serving well as 
a source of diagnostic features for the genus, the baculum does not seem to offer exceptional insight 
into suprageneric relationships. The large genus Pteropus, however, to some extent proves to be 
an exception, with various of its many species demonstrating considerable variation in bacular 
structure: on occasion differences between species equal those between some megachiropteran 
genera. 

Lanza ( 1 969) examined the baculum of Pteropus in detail and found that its bacular morphology 
did not conform to the classification proposed by Andersen (1912), a conclusion also reached by 
Davis ( 1 947) who examined only five species. Lanza found that in many cases bacula of an identical 
size and shape could be found among species belonging to the same group as well as to different 
groups; or that the baculum could be extremely different among forms apparently otherwise very 
closely related. Thus in this genus he found the baculum to be of limited value in phyletic analysis. 
Similarly, LaVal (1973ft) found that with one exception the bacula of the various species of the 
vespertilionid Rhogeessa are not sharply differentiated from each other: although in shape they 
show substantial geographic and individual variation within species they seem nevertheless to 
differ between species in areas of sympatry or near sympatry. Harrison & Brownlow (1978) found 
that individual variation in the baculum of adults of another vespertilionid, Scotophilus, was such 
that it rendered this structure of little or no value in species diagnosis in this difficult genus. 

Martin (1978) discussed the adaptive value of the baculum in bats, having found a wider range of 
structural variation among several pteropodid species than was previously thought. He considered 
that the baculum may have a number of roles of varying adaptive significance in supporting the 
penis, as a stimulatory structure, or in preventing urethral closure during the pressures of copula- 
tion. Although these might allow the baculum to maintain morphological stability within certain 
taxonomic units, this possible variability of function he thought tended to reduce its value in 
classification at the specific and subspecific levels. 

Despite these possible limitations, many authors admit at least the species-specificity of bacular 
variation among bats, using the baculum to provide additional characters to separate species that 
sometimes otherwise closely resemble each other. Some examples have been mentioned: others 
include Wallin (1969) who drew attention to bacular differences in Japanese Pipistrellus and who 
used such differences to define two species groups in Vesper tilio, or Baag0e (1973) who utilised 
bacular characters in comparing sibling species of European Myotis. Zubaid & Davison (in press) 
found the baculum to be specifically diagnostic among Malaysian Hipposideros. In some genera 
authors have routinely described and illustrated the baculum of new species: for instance Sinha 
(1969) in describing Pipistrellus peguensis compared its baculum with the bacula of the related 
species. Similarly, McKean et al. (1978) described and illustrated the baculum of 'Eptesicus' 
sagittula, comparing it with the bacula of other Australian 'Eptesicus', while Kitchener (1976) 
employed the baculum of ' Eptesicus' douglasorum in the same way. Bacular characters sometimes 
form an essential part of revisionary study, as by Kitchener et al. (1986) in defining and keying the 
Australo-Papuan representatives of Pipistrellus and Falsistrellus. The baculum has also featured 
in generic revision, Hill (1966a) for example describing and illustrating that of Philetor in the 
course of such a study, or (1976) that of the majority of the species of Hesperoptenus. 

Bacular variation has also been employed for generic and subgeneric distinction within the 



VESPERTILIONINE SYSTEMATICS 229 

Vespertilioninae. Wallin (1969) used penial characters in establishing Vespertilio as a genus distinct 
from Eptesicus and in recognising Hypsugo as a subgenus within Pipistrellus, while Topal (1970a) 
noted that bacular morphology allied the aberrant genus la more closely to Eptesicus than to 
Pipistrellus with which it had been associated by some authors. Heller & Volleth (1984) summar- 
ised published illustrations of the bacula of Pipistrellus, Eptesicus and some of their associated 
genera, drawing attention to their taxonomic implications. At a further systematic level, Pine et al. 
(1971) discussed the penial and bacular morphology of Antrozous and Bauerus in relation to the 
presumed affinities of these North American genera to the Australian and New Guinea genera 
Nyctophilus and Pharotis with which they have been associated in the subfamily Nyctophilinae. 

It is clear from the foregoing account that the baculum is regarded as a valuable source of 
diagnostic information by many students of chiropteran systematics. This seems especially true in 
the Vespertilioninae, a subfamily in which diagnosis and definition at both specific and generic 
levels is sometimes difficult if only the orthodox morphological characters of external, cranial and 
dental structure are to be relied upon. 



Nature and scope of this study 

The basis of the current classification of the Vespertilioninae was first set out in detail by Miller 
(1907), who recognised a total of thirty-two genera in the group, with diagnoses and short descrip- 
tive accounts. The classification of Miller was based chiefly on external, cranial and dental features. 
Tate (1942a) reviewed the characters used for diagnosis in some detail, dividing the subfamily 
into four main (tribal) aggregations, and attempting to quantify the interrelationships of its 
many genera. The major outlines of his classification have since been followed, sometimes with 
local modification as for instance by Koopman (19840, b, 1985) who subsumed the subfamily 
Nyctophilinae into the Vespertilioninae. Hill (1966) pointed out that the subfamily comprises a 
complex of closely interrelated genera separated in some instances by comparatively slender or 
even rather arbitrary distinctions, the patterns of relationship often obscured by parallelism or 
convergence. 

The narrowness of the orthodox distinctions that define many of the constituent genera of the 
Vespertilioninae has led to much taxonomic combination and recombination since Tate wrote. 
This situation is exemplified by the more extreme variants of classification that have been pro- 
posed. For example, Kuzyakin( 1944, 1950, 1 965) included Pipistrellus and Eptesicus in Vespertilio 
while Simpson (1945) included Glischropus, Scotozous, Nyctalus and la in Pipistrellus and Rhinop- 
terus, Hesperoptenus, Tylonycteris, Mimetillus, Philetor, Histiotus and Laephotis in Eptesicus. A yet 
more extreme viewpoint was adopted by Sokolov (1973) who considered that Vespertilio should 
include not only Pipistrellus and Eptesicus as was thought by Kuzyakin, but also all of the other 
above mentioned genera except Nyctalus. Horacek & Hanak (19850, b) commented that the 
concepts of Kuzyakin and Sokolov (with the inclusion of Nyctalus} might be provisionally 
accepted, at least until factual proof of paraphylly in the group was forthcoming. Nevertheless, 
they considered this to be a retrograde solution since it expresses nothing of the factual diversity of 
the group, proposing instead that the problematic taxa should be arranged in separate genera, their 
diagnoses then making their content clearer though narrower. Both Simpson and Sokolov also 
included Scotoecus and Scotomanes in Nycticeius as then understood, Baeodon in Rhogeessa, 
Glauconycteris in Chalinolobus and Dasypterus in Lasiurus to produce a heavily 'lumped' classifica- 
tion. The status of some such as Scotoecus, Dasypterus and Idionycteris has varied from one author 
to another for decades: in the Australian region Scoteanax and Scotorepens have recently achieved 
generic rank after many years as nominal subgenera (Kitchener & Caputi, 1985) while la has once 
again reverted to Pipistrellus (Koopman 1984a, b, 1985) after a brief spell with Eptesicus. The 
major variants of vespertilionine classification are summarised in Table 1. 

Many of the characters used to define taxa and relationships among the Vespertilioninae appear 
strongly adaptive and of equivocal value in generic and suprageneric systematics. Most concern 



230 J. E. HILL & D. L. HARRISON 

ear size and shape, tragal structure, the architecture of the skull, and the number and formation of 
the teeth. Zima & Horacek (1985) pointed out that the use of the morphological characters 
employed hitherto in the classification of the Vespertilionidae as a whole might not lead invariably 
to correct taxonomic conclusions, their degree of differentiation perhaps reflecting the orientation 
and intensity of selection pressure rather than actual phyletic relationships. These authors indi- 
cated an urgent need for new, sufficiently reliable and taxonomically useful criteria based on 
features that did not possess a direct adaptive significance, including among these the morphology 
of the reproductive organs and the baculum. 

Much weight has been placed in the past upon the progressive shortening of the muzzle apparent 
throughout the Vespertilioninae with concomitant reduction and loss of the incisors and pre- 
molars (Tate, 1942a). In the incisive dentition the first upper tooth (i 1 ) 3 is absent, as in all bats. 
Reduction results in the remaining inner tooth (i 2 ) becoming peg-like and unicuspid, although 
sometimes quite massive, in a reduction in size of the outer tooth (i 3 ), its displacement inwards 
or outwards, or in its eventual obsolescence or loss. In the mandible, the first (ij and second 
(i 2 ) incisor teeth are invariably present, but exceptionally the third (i 3 ) may be absent. The process 
also involves the reduction and loss of the second upper and lower premolars (pm}) and then 
of the anterior upper premolar (pm 2 ): thus the premolar formula ranges from pm rf y to pm 
5 i 5 f the first upper and lower premolars (pm}) being presumed to be those that are absent from all 
bats. Seven different combinations of incisors and premolars occur in the subfamily, if Antrozous 
and Bauerus are included. The full complement is usually taken as the primitive condition, the 
reduction and disappearance of teeth as derived. These are summarised in Table 2, which gives the 
incisive and premolar formulae usually attributed to each of the various genera. However, Myotis 
occur in which pm 3 or pmj are absent (Hill & Topal, 1973), thus in the first instance producing 
the formula typified by Lasionycteris or Plecotus, in the second the formula for Pipistrellus or 
Nyctalus; pm 2 may be absent from Pipistrellus to give the arrangement for Eptesicus, or may be 
present in Eptesicus to produce the formula for Pipistrellus (Hill & Topal, loc. cit.); i 3 is variable in 
Scotozous (of 45 examined, present in 34, absent from one side or the other in 8, completely absent 
from 3), when totally absent to produce the incisive formula that usually characterises Nycticeius 
and its associates (but Thomas & Wroughton ( 1 908) report an example of 'Nycticeius ' schlieffenii 
in which the left i 3 is present); pm 2 is variable in Scotoecus (Hill, 1974) and in Chalinolobus (Ryan, 
1966; Koopman, 1971), and very rarely may be present in 'Nycticeius' schlieffenii (Dobson, 1878; 
Thomas, 1890). 

Most genera of Vespertilioninae can be defined by other features besides those of the incisive and 
premolar dentition, although sometimes only in differing combinations. Thus although some 
species exist that combine the external features of Myotis with the dental formula of Pipistrellus to 
the extent that initially they (annectans, ridleyi) were described in the latter genus, other characters 
such as the form and structure of the tragus and the structure of the incisors enable them to be 
referred confidently to Myotis (Topal, 19706; Hill & Topal, 1973). Another (rosseti) was first 
described in Glischropus, subsequently removed to Pipistrellus by Hill (1969) and finally (with 
ridleyi) to Myotis by Hill & Topal (loc. cit.). However, the genera Pipistrellus and Eptesicus do not 
offer further conventional characters in this way and are separated for the most part by the 
presence of pm 2 in the former and its absence in the latter. Wallin (1969) and Hill & Topal (loc. cit.) 
discussed the variability of this tooth in Pipistrellus and Eptesicus in detail, the latter authors 
concluding that the presence or absence of pm 2 can have no universal validity in defining the two 
genera. Heller & Volleth (1984) also examined the relevance of pm 2 in separating Pipistrellus and 
Eptesicus and concluded that it does not seem to be a reliable characteristic, a classification based 
on it perhaps misrepresenting true relationship. Tate (\942a) recognised this difficulty but adhered 
to the conventional practice of separating the two genera by this feature, and indeed the majority of 
authors have retained the distinction as a matter of convenience, often using the extent of reduction 
and degree of displacement of pmf from the line of the toothrow as a diagnostic feature between the 
species of Pipistrellus. 



"Dental notation of Miller (1907) 



VESPERTILIONINE SYSTEMATICS 23 1 

Koopman (1975) has commented upon this problem. This author examined African species 
allocated variously to Pipistrellus and Eptesicus in an attempt to find some other character that 
would divide this large assemblage into two major groups. He could find none among the usual 
suite of morphological features. Although he found no African Pipistrellus species that closely 
resembled the hottentotus, tenuipinnis orfloweri groups of Eptesicus and no African member of the 
latter genus that was similar to thepipistrellus, Hesperus, savii or rueppellii groups of Pipistrellus as 
he defined them, he did find a resemblance amounting in some cases to virtual identity (if pm 2 was 
ignored) between the kuhlii group of Pipistrellus and the Eptesicus capensis group. Expanding a 
view first expressed by Tate (19420) in his account of Eptesicus, Koopman commented that it is 
probable that the anterior upper premolar has been lost more than once here, and that true phyletic 
relationships run across the 'generic' line. He thought that it is even possible that in some cases a 
'Pipistrellus species' and an 'Eptesicus species' are actually conspecific, but was of the opinion 
that the available material was insufficient to establish this with certainty for any such pair at 
the present time. Although retaining Pipistrellus and Eptesicus as separate genera since he 
believed that the problem should be attacked on a cosmopolitan basis, he remarked that such an 
arrangement is almost certainly wrong. More recently, Horacek & Hanak (1985-1986) have 
offered further definitions of Pipistrellus, Hypsugo and Eptesicus. 

Many varying interpretations can be placed upon external, cranial and dental morphology or on 
karyological data in the Vespertilioninae. These range from the relationship of one species to 
another to suprageneric relationships, even to the view that Pipistrellus and Eptesicus may be 
polyphyletic. Menu (1984), for example, remarked that an exhaustive odontological study of 
the Vespertilioninae indicated that Pipistrellus includes species wrongly associated by reason 
of their identical dental formula, but which are not related. Williams & Mares (1978) discussed the 
karyology of Eptesicus, which as currently defined they thought seemed to be a composite taxon, 
encompassing perhaps several phyletic lines of pipistrelloid species with reduced numbers of 
premolars. Heller & Volleth (1984) suggested that Pipistrellus may be a very heterogeneous 
assemblage and after reviewing the relevance of pm 2 in separating this genus from Eptesicus 
considered the baculum to be a more reliable guide to the phylogeny of the species of Pipistrellus 
and Eptesicus, using its features to supplement their findings from karyological data. Many years 
before this Tate (19420) remarked that it seemed probable that study of the baculum in the 
Microchiroptera would yield valuable results, with the implication that this might have signifi- 
cance in the classification of the Vespertilionidae. Indeed, Tate records that G. M. Allen had 
gathered together a number of bacula representing many of the species of Pipistrellus which he 
intended to employ in revising the genus. Moreover, relatively few species of the nominal genera 
Pipistrellus and Eptesicus have so far been studied: the impression gained from the literature is 
that Pipistrellus as currently understood is dignified chiefly by a long, slender shafted baculum and 
most Eptesicus as it is presently classified by a small, triangular structure, which we have found not 
to be the case. 

Initially our intention was to examine the bacula of as many species of Pipistrellus as possible to 
establish the pattern of bacular variation within the genus, and to compare it with the species 
groupings proposed by Tate (19420) and by Koopman (1973, 1975). As the work progressed, 
however, it became increasingly apparent that its implications extended far beyond the limits of 
this nominal genus and that it was necessary in addition to study the bacula of Eptesicus so far as we 
were able, and to examine the boundary between these two conventional groupings. Finally, to 
place our findings in proper perspective, we have surveyed the bacula of most of the remaining 
genera of the Vespertilioninae and have attempted to assess the generic significance of bacular 
variation in the subfamily. We have also examined the bacula of Nyctophilus and Pharotis, both 
usually referred to the closely related subfamily Nyctophilinae. This has been united recently with 
the Vespertilioninae (Koopman, 19840, b, 1985) and is thus relevant to our study. 

We have made no detailed examination of the gross morphology of the chiropteran penis except 
insofar as it is reflected by bacular structures. Nor have we attempted to study its histomorphology. 
These features are discussed by Smith & Madkour (1980) in an effort to elucidate their relevance to 
interordinal and infraordinal phylogenetic relationships, and who review earlier studies of penial 
morphology. 



232 J. E. HILL & D. L. HARRISON 

Materials and methods 

We have been able to examine bacula from the majority of species currently listed in Pipistrellus 
and from most of those presently assigned to Eptesicus. In a few instances we have relied 
upon illustrations and descriptions from the literature. Similarly, for the other genera of the 
Vespertilioninae our study material has been drawn chiefly from specimens and to a much lesser 
extent from the published works of others. The specimens that we have examined are listed in 
Appendix 1 . Our aim as far as genera other than Pipistrellus and Eptesicus are concerned has been 
to provide illustrations of representative bacula, but in those instances where bacular structure has 
not before been studied we have endeavoured to examine as many species within each genus as the 
available specimens permitted. Clearly, the material available to us has been quite inadequate to 
establish the extent of individual variation in any one species or subspecies. While only adult 
specimens (wing epiphyses fully fused) have been used, we have necessarily had to accept that for 
the majority of species our data is limited. We have concentrated therefore on studying and 
comparing the gross morphology (size, shape, gross structure) of the bacula that we have examined 
in an attempt to identify similarities, differences and general trends. The finer details perhaps more 
valuable in species distinction have received much less attention, although where it is known that 
species are difficult to separate by conventional means attention has been drawn to bacular features 
that may assist in identification. 

The specimens used in this study have been drawn almost entirely from the collections of the 
British Museum (Natural History), London (BM(NH)) and the Harrison Zoological Museum, 
Sevenoaks, Kent (HZM). Apart from these we have been able to examine one from the Natur- 
historisches Museum, Wien (NMW), by courtesy of Dr K. Bauer, and one from the Carnegie 
Museum of Natural History, Pittsburgh (CMNH), an anomalous specimen loaned for identifica- 
tion by Dr D. A. Schlitter, while Dr K. F. Koopman generously brought to London an example of 
Nycticeius humeralis from the American Museum of Natural History, New York (AMNH) from 
which a much needed baculum was obtained. Specimens prepared many years ago at the British 
Museum (Natural History) are dry, sometimes mounted on card: the remainder have been pre- 
pared in the course of this study. This has been accomplished by maceration for a short period in a 
5% solution of potassium hydroxide to which a small quantity of alizarin red has been added, after 
which the grosser macerated tissue was removed by dissection, the specimen then being cleared and 
stored in glycerin. 

Drawings have been prepared using either a stereoscopic microscope with graticule scale and 
attached camera lucida, or freehand using a similar instrument. A few were drawn freehand using a 
stereo projection microscope with travelling micrometer stage. The wide range of size variation 
among vespertilionine bacula (for example from a length of 1 mm or less to as much as 9 or 10 mm 
in Pipistrellus) has necessitated the use of several scales of magnification. So far as possible all 
drawings on any one page of figures are at the same magnification, with an appropriate scale: to 
facilitate comparison the varying magnifications used follow an arithmetic progression whereby 
each successive larger value is twice its predecessor. It has not always been possible to conform to 
this arrangement, especially where drawings have been prepared from published illustrations. As a 
rule dorsal (D) and right lateral (RL) views of each baculum are provided: rarely through damage 
the left lateral (LL) aspect is given. Occasionally where it is of especial interest a half ventral (RVL 
or LVL) drawing has been made, and in a few instances where drawings have been taken from the 
literature it has been necessary to give the ventral (V) rather than the dorsal aspect. 

Authorship and responsibility 

We take joint responsibility for the results and opinions put forward and expressed in this paper, 
and for the new names proposed therein. 

The baculum of Pipistrellus 

Four bacular types have been identified within the nominal genus Pipistrellus. With some excep- 
tions, modification and combinations, these are in broad agreement with the groupings of species 



VESPERTILIONINE SYSTEMATICS 233 

proposed by Tate (19420) and Koopman (1973, 1975). The classifications of Tate and Koopman 
are summarised in Table 3. 

(1) An elongate structure (Fig. la) with a slender shaft and paired basal flanges (e.g. Figs 2a-c, 
3, 4, 5), the ventral surface of the proximal part of the shaft transversely concave, its distal part 
cylindrical or nearly so; in profile the base in line with the shaft or more or less deflected downward 
at an angle to it; the shaft may be more or less straight, flexed or variously curved in the vertical 
plane, while the tip is generally bifid or forked and may be directed ventrally to a greater or lesser 
extent. 

Species aggregations in which this type of baculum is found include the abramus, pipistrellus, 
coromandra and tennis groups of Tate (19420); Koopman (1973) amalgamated these to form a 
pipistrellus group to which he added (1975) the African nanus and permixtus. However, nanus 
(Fig. 6b) proves to have a very different baculum, as does imbricatus (Fig. 9a), included by Tate in 
the coromandra group and thus by Koopman ( 1 973) in the pipistrellus group. Pipistrellus babu (Fig. 
4a), provisionally placed by Tate in the kuhlii group, also has the long, relatively straight baculum 
characteristic of this part of the division, as do endoi (Imaizumi, 1959) (Fig. 3b) and peguensis 
(Sinha, 1969) (Fig. 15c), both described since Tate wrote. The more recently described westralis 
(Koopman, 1984c) (Fig. lOd), adamsi (Fig. lOc) and wattsi (Fig. lOg) (Kitchener et al, 1986) also 
belong with tenuis (Fig. 9d) and its allies in this grouping. Taxa referred to the ceylonicus group by 
both Tate and Koopman (1973) prove to have this bacular structure, as do those that have been 
examined of the rueppellii group (Figs 7e, f, lOa, b) of Koopman (1975). Pipistrellus kuhlii and its 
associates (Figs 5a-d, 6c) also belong in this division. In these, however, the basal lobes of the 
baculum are sharply angled to the shaft in the vertical plane, and this bacular profile is very 
characteristic of kuhlii and its relatives. The shaft is straight, without flexion, and the tip is usually 
bifid and not directed ventrally. Koopman (1975) included anchietae (Fig. 6e) in the kuhlii group, 
but this proves to have a very different bacular configuration. 

A long-shafted baculum of this type occurs with little modification in the majority of the 
Australian species (Figs lla-f, 12k) currently referred to Eptesicus, in Nyc talus (Fig. lOf), in 
Scotozous (Fig. 16d) (to which rueppellii and its immediate associates have sometimes been 
referred) and in Scotoecus (Fig. 20a-e), in which the 'horns' of the bifid tip extend in some instances 
almost to form a ring, a condition foreshadowed in Pipistrellus pater culus (Fig. 3c). The Australian 
Scoteanax (Fig. 16i) and Scotorepens (Figs 16g, h, 21e, also share this bacular type: in Scoteanax 
the 'horns' at the tip have become a transverse bar, but the species of Scotorepens retain the bifid 
or slightly bifid tip. A similar long-shafted baculum but with a simple tip occurs in the genera 
Hesperoptenus (Fig. 21a-c, g) and Chalinolobus (Fig. 17a-e). The baculum of Glischropus (Fig. 
18a), although very small, is also of this type, with paired basal lobes, a slender shaft, and bifid tip. 

(2) A very small structure (Fig. Ib), consisting of a broad base with two basal lobes (e.g. Figs 2d, 
e, 9c, h), supporting a short, very slightly hollowed shaft. This bacular type is found in subflavus 
(Fig. 2d), circumdatus (Fig. 2e), societatis (Fig. 9c) and the more recently described cuprosus (Hill & 
Francis, 1984) (Fig. 9h). 

(3) A relatively short, stout shafted baculum (Fig. Ic), sometimes with expanded base and tip 
(e.g. Figs 6a, b, 7a, h, 8e, f), the base on occasion divided into paired lobes, sometimes angled 
vertically to the line of the shaft, which is fluted ventrally rather than mostly cylindrical; tip when 
expanded having its anterior edge sometimes divided into several irregular serrations and on 
occasion downwardly directed. 

Such bacula are found in the savii group of Tate (19420) and Koopman (1973, 1975) but not in 
maderensis (Fig. 5b) which was put into the savii group by both authors. Its baculum is however 
quite different and is like that of kuhlii and its associates. Pipistrellus anchietae (Fig. 6e), referred to 
the kuhlii group by Koopman (1975) also belongs with savii, and the same bacular type occurs in 
nanus (Fig. 6b), allocated with permixtus to the pipistrellus group by the same author. We have been 
unable to examine the baculum of permixtus but that of nanus and ofhelios (Fig. 6d) is of the type 
characteristic of this division, with its basal part quite sharply flexed to the shaft although not 
especially deep, and with an expanded, downwardly directed distal part. Pipistrellus eisentrauti 



234 J. E. HILL & D. L. HARRISON 

(Fig. 9g), referred to the rueppellii group by Koopman (1975) also shares this bacular type. The 
North American Hesperus (Fig. 8d) was placed by this author in a Hesperus group, with the African 
musciculus. It has a robust baculum of the type found in this division, somewhat flattened, without 
basal lobes but broadened just beyond the base, the shaft narrowing towards the tip; the ventral 
surface is shallowly fluted throughout its length. Unfortunately, no baculum has been available for 
musciculus but provisionally it is referred to this division on other grounds. 

This bacular class also includes imbricatus (Fig. 9a), referred to the coromandra group by Tate 
(1942a) and by Koopman (1973), and pulveratus (Fig. 8c), lophurus (Fig. 8f), and kitcheneri (Fig. 
8e) which Tate placed in the qffinis group (vide infra). Koopman (loc. cit.) followed this lead with 
respect to kitcheneri but did not include pulveratus and lophurus since these were extralimital to 
his study. Pipistrellus bodenheimeri (Fig. 9f), described (Harrison, 1960) since Tate wrote and 
extralimital to Koopman (loc. cit., 1975) also belongs in this group. Tate referred macrotis, vorder- 
manni and curtatus to the savii group on account of their reduced pm 2 but indicated that this 
allocation might not be tenable: however, the baculum of macrotis shows that it should be placed in 
this group. This author also created the joffrei group to include joffrei, anthonyi, brachypterus, and 
stenopterus: brachypterus has since proved to be a Philetor (Hill, 1971). We have been unable to 
examine the baculum in either joffrei or anthonyi but that of stenopterus (Fig. 7h) indicates that it 
belongs here. Both joffrei and stenopterus have been referred variously to Nyctalus (Chasen, 1940; 
Ellerman & Morrison-Scott, 1951) or to Pipistrellus (Tate, 19420; Hill, 19660) but the baculum of 
stenopterus has no resemblance to the long-shafted structure of the former genus. 

(4) A relatively large, short but strong baculum (Fig. Id), broad, with little or no proximal or 
distal expansion (e.g. Figs 8a, b, g, lOh), the ventral surface transversely deeply concave so that it is 
strongly arched or fluted throughout its length. This grouping includes affinis (Fig. 8a) andpetersi 
(Fig. 8b), placed in the qffinis group by Tate (19420) and in the case ofpetersi in the same group by 
Koopman (1973), qffinis being extralimital to his study, together with the Australian tasmaniensis 
(Fig. 8g) for which Tate maintained a tasmaniensis group. A further Australian form, mackenziei 
(Kitchener et al., 1986) (Fig. lOh) is very like tasmaniensis and also belongs here. To some extent 
this grouping is linked to the previous division by pulveratus, imbricatus, lophurus, kitcheneri and 
their immediate associates: Tate allocated all except imbricatus to the affinis group. 

The baculum of Eptesicm 

We have been able to identify three bacular types among the species currently referred to the 
nominal genus Eptesicus. There is no single reference for species groupings in this aggregation of 
species, but for African forms these bacular types agree almost exactly with the species groups 
defined by Koopman (1975). 

(1) A more or less triangular structure (Fig. le), its apex occasionally drawn out into a slight, 
short shaft, the base widened and sometimes slightly lobed but the tip not expanded, usually more 
or less pointed or gently rounded (e.g. Figs 13, 14a, c). This type of baculum is flattened, with little 
ventral fluting or concavity: there is little vertical flexion either of the base or of the more distal part, 
and the tip is not deflected downwards. So far as we have been able to establish, this bacular class 
occurs in all of the forms that are currently referred to Eptesicus from the New World, Europe and 
Asia, and in the African forms that Koopman (1975) included in the serotinus andfloweri groups. 

(2) The structure in a small group of species, wholly Australian, in which the baculum has 
usually a long cylindrical or slightly fluted shaft with paired expanded basal lobes (Fig. If) and 
usually a blunt tip (e.g. Fig. 1 la-e), very similar in fact to the first of the bacular types that we have 
described for Pipistrellus. This grouping includes pumilus (Figs 1 la, b, 12k), vulturnus (Fig. 1 le), 
douglasorum (Fig. 1 Id) and regulus (Fig. 1 le): in pumilus and douglasorum the shaft and base may 
be flexed rather like those of Pipistrellus kuhlii and its immediate relatives, while in regulus the shaft 
has an expansion just behind the tip (McKean et al., 1978; Kitchener, 1976). A further species, 
sagittula (Fig. llf), also appears to belong here, its baculum being perhaps a shorter-shafted 
version of this type. 



VESPERTILIONINE SYSTEMATICS 235 

(3) A slender-shafted baculum (Fig. Ig), usually with distinct paired basal lobes, sometimes 
angled or flexed to the line of the shaft, which is cylindrical, with variously expanded tip, the distal 
expansion varying from a downwardly directed spatulate plate to a large, anteriorly directed, 
downwardly deflected lobed structure (e.g. Figs 12a-j, 14b). Taxa with this type of baculum are 
wholly African and the grouping comprises those forms referred to the capensis and tenuipinnis 
group by Koopman (1975). 



The baculum in other Vespertilioninae 

Before considering the implications of bacular morphology in relation to the systematics of 
Pipistrellus and Eptesicus, a brief review of bacular types in the remaining genera of the 
Vespertilioninae will serve to place these nominal genera in the perspective of bacular structure in 
the subfamily as a whole. 

My otis (Fig. 19i, j). The baculum of Myotis has been figured and described by numerous 
authors. Palaearctic species have been studied by Topal (1958), Hanak (1965, 1970, 1971), Wallin 
(1969), Atallah (1970) and Baag0e (1973), among others. Nearctic and Neotropical species have 
been examined by Hamilton (1949), Wimsatt & Kallen (1952), Krutzsch & Vaughan (1955), Davis 
& Rippy (1968), Genoways & Jones (1969), LaVal (19730) and Warner (1982). In this genus the 
baculum is much like a small saddle. In profile the base and tip are slightly elevated, the baculum 
ventrally slightly concave. In dorsal aspect the baculum is more or less triangular or projectile- 
shaped, anteriorly bluntly or sometimes more sharply pointed, the base divided to a greater or 
lesser extent into two lobes, the ventral surface deeply fluted. In some instances at least there is 
evidently distinctive variation between species and it is possible that some clear infrageneric 
division into bacular types might be made. There is also apparently considerable individual varia- 
tion in some species, leading LaVal (19730) in the case ofnigricans to suggest the possibility that 
the material that he studied was a composite of sibling species. However, for so large a genus 
there seems to be a surprising degree of broad homogeneity in gross bacular structure. We have 
examined relatively few bacula from such a well known genus, but it is of some interest to note that 
the baculum ofridleyi (Fig. 1 9i), described and retained for many years in Pipistrellus on account of 
its dentition which corresponds to that genus (Hill & Topal, 1973) is undeniably of the type 
characteristic of Myotis. 

Pizonyx (Fig. 19k). Figured and described by Hamilton (1949). The baculum is similar to that of 
Myotis, but lacks much of the saddle-like appearance, and is more triangular in dorsal aspect, 
tapering distally to a flattened, slightly elevated tip; it is also elevated proximally so that in profile 
the base inclines slightly upwards. Ventrally the baculum is slightly fluted. 

Lasionycteris (Fig. 17f). Figured and described by Hamilton (1949). The baculum has a large, 
swollen, bilobed base, a long cylindrical shaft, and a slight distal enlargement, the tip and base 
elevated dorsally. Proximally, there is sometimes a flattened dorsal prominence on the base, its 
bilobed extremity projecting beyond the main bulbous part. 

Plecotus (including Corynorhinus). The Old World forms (Plecotus, Figs 14d, 19g, h) are figured 
and described by Topal (1958), Lanza (1960) and Ibanez & Fernandez (1986), American taxa 
(Corynorhinus, Fig. 15f-h) by Nader & Hoffmeister (1983). In most the baculum is arrow-head- 
shaped, slightly saddle-like, with basal lobes and broad, short distal part, the base elevated 
dorsally, the ventral surface deeply fluted: in two taxa (auritus, Fig. 19g, teneriffae, Fig. 14d) it has a 
longer, more slender shaft with paired basal lobes and is less saddle-like. 

Idionycteris (Fig. 15e). Figured and described by Nader & Hoffmeister (1983). An elongate 
baculum, with triangular basal plate, its apex directed posteriorly, and narrow shaft, curved 
dorsally and ventrally shallowly grooved. 

Euderma. The baculum in this genus is so far apparently unknown. 



236 J. E. HILL & D. L. HARRISON 

Barbastella (Fig. 18j). Figured and described by Topal (1958). A small, saddle-like baculum 
similar in many ways to that of Plecotus, with elevated base, narrowed distal part which is 
upwardly curved, and with slightly raised, elevated tip. 

Rhogeessa (Fig. 18k). Figured and described by LaVal ( 1 9736). A small baculum with expanded, 
bilobed base and short, stubby shaft lacking any distal modification, the shaft ventrally fluted. 

Baeodon (Fig. 15b). Figured and described by Brown et al. (1971) and LaVal (19736). Baculum 
very like that of Rhogeessa but with shorter shaft. 

Nycticeius. Australian forms hitherto referred to Nycticeius have been recognised as Scoteanax 
and Scotorepens by Kitchener & Caputi (1985). As therefore it is currently understood, Nycticeius 
includes two species, humeralis from North America and schlieffenii from Africa. These have 
widely differing bacula. In humeralis (Fig. 1 7k) the baculum is blade-like, with short, narrow shaft, 
the base thickened, proximally forming a prominent angle which inclines towards the ventral 
surface, the distal portion deep, with convex walls which terminate in an ascending point. This 
structure is figured by Hamilton (1949) who remarked that it differed markedly from the bacula of 
other [North American] genera: in fact it is not closely approached by any other vespertilionine. 
The baculum of schlieffenii (Fig. 16e), by contrast, has a broad bilobed base with tapering, fluted 
shaft, its tip unmodified, bluntly pointed, and unexpanded. Moreover, the bacular morphology of 
the Australian species formerly referred to Nycticeius supports their separation from this genus. 
This matter is discussed more fully below. 

Otonycteris (Fig. 16a). Figured and described by Wassif & Madkour (1972), Fairon (1980) and 
Wassif, Madkour & Soliman (1984). An unusual baculum, mostly a more or less parallel-sided 
narrow shaft, the base and tip not expanded, both strongly elevated dorsally, the shaft tapering 
distally to a raised tip. 

Lasiurus (Fig. 191). Figured and described by Hamilton (1949). A slipper-like baculum with 
broad, dorsally elevated base, a short shaft, fluted ventrally, and with slightly expanded and 
elevated tip. 

Dasypterus (Fig. 18f). Figured and described by Brown et al. (1971) and Hamilton (1949). 
Baculum like that of Lasiurus but tip as a rule not upturned. 

Antrozous (Fig. 18b). Figured and described by Krutzsch & Vaughan (1955) and Pine et al. 
(1971). Baculum broadly triangular in dorsal view, tapering to broad, blunt point, fluted ventrally, 
and with the base elevated dorsally. It is very different from the baculum in Nyctophilus and 
Pharotis, and from that of Otonycteris, with which genera Antrozous has been associated in the 
past. 

Bauerus (Fig. 15i). Figured and described by Pine et al. (1971). Baculum like that of Antrozous 
but smaller and narrower, the distal part not upcurved. 

Scotomanes (Fig. 18g). A short baculum with broad, bilobed base merging into a very narrow, 
short cylindrical shaft with no distal expansion, lacking any upward deflection either proximally or 
distally. 

Scotophilus (Fig. 17g-j). Figured and described by Harrison & Brownlow (1978). Baculum 
irregularly sub-rectangular or sub-triangular, flattened, anteriorly usually bluntly rounded, 
slightly concave in ventral transverse section, with slight basal lobes. 

Vespertilio. Figured and described by Topal (1958) and Wallin (1969). In two species (murinus 
and orientalis) the baculum is situated at the base of the penis, which is supported by a cartilaginous 
pseudobaculum. In the third (superans) the baculum is situated not at the base of the penis but 
midway along the shaft, and there is no pseudobaculum. The baculum in orientalis (Fig. 21j) and 
superans is flattened and triangular, with a broad, bilobed base, tapering anteriorly to a narrow 
point and with slight vertical flexion. The baculum of murinus (Fig. 21i) is broad but less triangular 
in outline, and has a wide, bluntly rounded distal part. The bacula of orientalis and superans in 
particular are similar in many respects to those of the Eptesicus serotinus group (vide supra). 



VESPERTILIONINE SYSTEMATICS 237 

Histiotus (Fig. 18c-e). A very small baculum, with expanded bilobed base and short, narrow 
cylindrical shaft, its tip unexpanded, the base and tip deflected slightly upwards. 

la (Fig. 2 Id). Figured and described by Topal (19700). A large, flattened, triangular baculum 
similar to those of the Eptesicus serotinus group (vide supra). 

Tylonycteris (Fig. 18h, i). Baculum small, similar to that of Histiotus or to those of the Eptesicus 
serotinus group (vide supra), but with the distal part extended into a narrowed shaft and with 
relatively wider, expanded base with a slight trace of basal lobes. 

Glauconycteris (Fig. 19a-f). Baculum very small and somewhat variable within the genus, but 
mostly more or less triangular, with some modification, usually as reduction, to a deeply lobed base 
with a short, blunt distal portion. However, on occasion the base is slightly or considerably 
expanded and the distal portion lengthened to a short shaft. 

Mimetillus. We have been unable to establish the presence of a baculum in this monotypic genus, 
from which it appears to be lacking. 

Eudiscopus. The baculum ofEudiscopus (if present) is apparently unknown. 

Nyctalus (Fig. 100- Figured and described by Topal (1958) and Lanza (1959). A long, slender 
baculum with narrow basal lobes, a long cylindrical shaft, and slightly bifurcated tip. 

Laephotis (Fig. 160- Baculum with expanded, bilobed base, narrow fluted shaft and broadly 
expanded tip with slight downward deflection, a small protuberance on its upper surface. Similar in 
many respects to the baculum in the Eptesicus capensis and E. tenuipinnis groups (vide supra). 

Glischropus (Fig. 18a). A very small baculum, with paired basal lobes, narrow cylindrical shaft 
and slightly expanded, bifid tip. 

Scotozous (Fig. 16d). Figured and described by Sinha (1976). A long baculum with slight basal 
lobes, a narrow, fluted shaft, and slightly bifid tip, the shaft slightly flexed. 

Scoteanax (Fig. 1 6i). Figured and described by Kitchener & Caputi ( 1 985). A long baculum with 
strong, expanded bilobed base, a slender cylindrical shaft, and with the tip embellished into a short, 
transverse bar. 

Scotorepens (Figs 16g, h, 21e, f)- Figured and described by Kitchener & Caputi (1985). A long 
baculum with expanded, bilobed base and slender, cylindrical shaft, the tip slightly expanded and 
bifid, the 'horns' deflected ventrally. 

Scotoecus (Figs 20a-e, 21h). A long, slender baculum with slightly expanded and bilobed 
base, long cylindrical shaft and an expanded, bifurcated tip, the 'horns' extending ventrally and 
sometimes curving to form an almost complete ring. 

Philetor (Fig. 16b). Figured and described by Hill (19660). A strong but relatively short baculum 
with paired basal lobes, a short, fluted shaft, and expanded rugose tip, the base and tip elevated and 
deflected upwards. 

Hesperoptenus (Fig. 21a-c, g). Figured and described by Hill (1976) and Hill & Francis 
(1984). Baculum long and slender, with paired basal lobes, a flattened, ventrally fluted shaft, and 
unmodified, rounded tip. 

Chalinolobus (Fig. 17a-e). Baculum long, with clearly defined basal lobes, a long cylindrical 
shaft, and an expanded tip, the expansion sometimes slight, considerable, or bifid with two obtuse 
projections. 

Although the two genera are commonly referred to a separate subfamily, the Nyctophilinae, for 
purposes of comparison we have also examined the baculum in Nyctophilus and Pharotis. 

Nyctophilus (Figs 16c, 22a-g). Figured by Churchill et al. (1984). A long, rather broad baculum 
with scarcely expanded, bilobed base and a broad shaft tapering distally to a blunt point, or wider 



238 J. E. HILL & D. L. HARRISON 

terminally with a median emargination to produce a shallowly bifid tip; shaft ventrally deeply 
fluted. 

Pharotis (Fig. 22h). Baculum similar to that of Nyctophilus but shaft narrower, tapering to 
slightly expanded tip. 



Systematic considerations 

The majority of genera in the Vespertilioninae have bacula which overall display a wide range of 
variation in their gross morphology. Most have a distinctive baculum: where closely similar bacula 
occur in genera currently recognised as distinct, as for example in Scotozous and Pipistrellus , then 
close relationship has been presumed on other morphological grounds. We are thus persuaded that 
in this subfamily the baculum can be used as a guide to infrageneric and intergeneric classification, 
although it seems that its value as a suprageneric indicator may be less. In the same way, although 
we have not explored the point in detail, it has become apparent that in many genera the minor 
details of bacular morphology can be used to assist in species distinction. These considerations 
have led us to the view that the very dissimilar bacular types that we have been able to identify and 
define within Pipistrellus and Eptesicus do indeed reflect natural groupings and show that Eptesicus 
as it is currently defined is a composite. Certainly it seems true to say that the current classifi- 
cation of both nominal genera does not properly reflect the relationships that we believe bacular 
morphology suggests exist within and between them. 

Genus Pipistrellus Kaup, 1829 

Pipistrellus Kaup, 1829:98. Vesper tilio pipistrellus Schreber. 

Romicia Gray, 1838: 495. Romicia calcarata Gray = Vespertilio kuhlii Kuhl. 

Rotnicius Blyth, 1840: 75. Variant of Romicia Gray. 

Hypsugo Kolenati, 1956: 131. Included Vespertilio maurus Blasius = Vespertilio savii Bonaparte, and 

Vespertilio krascheninnikowii Eversmann. Type species fixed as Vespertilio savii Bonaparte by Wallin 

(1969). Valid as a subgenus. 
Nannugo Kolenati, 1856: 131. Included Vespertilio nathusii Keyserling & Blasius, Vespertilio kuhlii Kuhl 

and Vespertilio pipistrellus Schreber. 
Alobus Peters, 1868: 707. Vespertilio temminckii Cretzschmar = Vespertilio ruppellii Fischer. Preoccupied by 

Alobus Le Conte, 1856 (Coleoptera). 

Euvesperugo Acloque, 1899: 35. Included six species, one being Vespertilio pipistrellus Schreber. 
Eptesicops Roberts, 1926: 245. Scotophilus rusticus Tomes. 
Neoromicia Roberts, 1926: 245. Eptesicus zuluensis Roberts. Valid as a subgenus. 
Vansonia Roberts, 1946: 304. Pipistrellus vernayi Roberts = Vespertilio ruppellii Fischer. 
Vespadelus Iredale & Troughton, 1934: iii, 95. Scotophilus pumilus Gray. Nomen nudum. 
Vespadelus Troughton, 1943: 348. Scotophilus pumilus Gray. Valid as a subgenus. 
Registrellus Troughton, 1943: 349. Pipistrellus regulus Thomas (see Hill, 19666). 
Falsistrellus Troughton, 1943: 349. Vespertilio tasmaniensis Gould. Valid as a subgenus. 
Perimyotis Menu, 1984: 409, 415. Vespertilio subflavus F. Cuvier. Valid as a subgenus. 
Parastrellus Horacek & Hanak, 1985a: unpaginated; 19856: 62; 1985-1986: 15, fig. 4. Pipistrellus Hesperus 

H. Allen. Nomen nudum. 

The genus Pipistrellus cannot be diagnosed by conventional morphological characters that are 
individually exclusive. Its current definition rests on Miller (1907) who based his diagnosis on the 
structure of i 2 which is simple or has a well developed secondary cusp; on the reduction of i 3 which 
is smaller than i 2 but nevertheless extends beyond the cingulum of that tooth; on rather short 
canines, c 1 often but not invariably with incipient secondary cusp on its posterior edge; and on the 
absence of pmf to give the dental formula iff |, c}, pm^f |, m} 2 ^ = 34, with pm 2 barely or not in the 
toothrow. He remarked that the members of the genus were recognisable by their dental formula, 
large i 3 , unmodified skull and ears, and the normally long fifth finger. 

The definition of the genus is briefly discussed by Tate (1942a), Ellerman & Morrison-Scott 
(1951) and Kitchener et al. (1986). All recognised the unreliability of the presence or absence of 
pm 2 as a prime diagnostic character, Ellerman & Morrison-Scott also remarking that 'strictly 



VESPERTILIONINE SYSTEMATICS 239 

speaking Pipistrellus is not more than a subgenus of Eptesicus, which itself might be referred to 
Vespertilio', but for convenience they and most other recent authors have followed the conven- 
tional distinction. It is clear from the foregoing account of the baculum in the Vespertilioninae that 
the species allocated to Pipistrellus can be separated from most other vespertilionine genera by 
their bacular morphology: those genera which have bacula similar to those of some Pipistrellus 
species (e.g. Nyctalus, Scotozous) can be defined by other morphological features of the skull and 
dentition, as they were by Miller (loc. cit.). 

Species groups in Pipistrellus are difficult and in some instances almost impossible to define on 
external, cranial and dental characters: most (Tate, 19420; Koopman, 1973, 1975) are brought 
together by combinations of characters with few or sometimes no exclusive features. Some species 
of Pipistrellus, moreover, appear difficult to separate from some of Eptesicus (Koopman, 1975; 
Heller & Volleth, 1984) except by the presence or absence of pm 2 which is itself evanescent. Bacular 
morphology appears to offer at least a partial solution to this difficulty, at the same time indicating 
that the genus as currently understood is a composite of several different groups of species, as 
suggested by Heller & Volleth (loc. cit.) on the basis of its known karyology. 

Chromosomal features so far as they have been established in Pipistrellus are reviewed by Heller 
& Volleth (1984) and Zima & Horacek (1985). Their summaries demonstrate that karyologically 
Pipistrellus as currently constituted is a very heterogeneous and diverse group, with 2N varying 
from 26-44 and FN from 44-60. At this stage we have been unable to find any consistent correla- 
tion between the chromosome formulae that these authors quote for various species and the 
groupings that we recognise on bacular and other grounds. However, many species remain to 
be studied karyologically and it appears from Zima & Horacek that for the present karyotype 
variability in the Vespertilioninae may be only of limited value as a taxonomic criterion. 

While at present we would not support the generic division of Pipistrellus as has been indicated 
or suggested by Menu (1984), Horacek & Hanak (19850, b} or Kitchener et al. (1986) since besides 
bacular features there appear to be few or no characters reported for its constituent groups that 
would support this wider separation, we consider that the divisions apparent within the genus 
justify subgeneric recognition. Wallin (1969) has already anticipated this view to some extent, 
employing Hypsugo Kolenati, 1856 for P. savii, in part on bacular grounds. Horacek & Hanak 
(1985-1986) recognised Hypsugo as a distinct genus. The gross morphology of the baculum 
also indicates that the Australian taxa formerly referred to Eptesicus should be transferred to 
Pipistrellus, as Heller & Volleth (1984) suggested, and that the African forms hitherto allocated to 
the capensis and tenuipinnis groups of Eptesicus also represent Pipistrellus as these authors inferred 
on account of their known karyology. Thus we would classify Pipistrellus in the following manner, 
listing included taxa without distinction as to taxonomic rank: some are not necessarily valid 
species or subspecies and for obvious reasons we have been unable to examine every named form in 
the genus. 

Subgenus Pipistrellus (Pipistrellus) 

Baculum long, with strong, extended shaft, well developed basal lobes, nearly always with a bifid 
tip. Braincase high, rounded, not flattened, sometimes globose; postorbital region usually wide; 
cranial profile generally straight or nearly straight from occiput to nares; interdental palate longer 
than wide; maxillary toothrows parallel or only slightly convergent anteriorly; i 2 generally 
bicuspid; pm 2 usually large, pm 2 not greatly reduced, usually about 3/4 crown area of pm 4 . 
Pipistrellus kuhlii and its associates differ slightly in almost unicuspid i 2 ; greatly reduced i 3 , much 
reduced pm 2 and more reduced pm 2 although some of these features occur in isolation in other 
species of the subgenus. 

(a) pipistrellus group 

Basal lobes of baculum more or less in line with the bacular shaft in the vertical plane; i 2 bicuspid, 

but not strongly so, cusps not deeply divided. 

(a) (i) pipistrellus subgroup. Braincase high, rounded; postorbital region wide; supraorbital 
region not widened or swollen; rostrum long, not greatly broadened, with shallow median rostral 



240 J. E. HILL & D. L. HARRISON 

depression; cranial profile almost straight from occiput to nares, slightly depressed over anterior 
part of orbit; premaxillae not shortened; zygomata slender, lacking any jugal eminence; interdental 
palate longer than wide; maxillary toothrows parallel for most of their length, anteriorly slightly 
convergent; short bony post-palate; slight basial pits; i 2 bicuspid, posterior cusp 1/2-3/4 the height 
of anterior cusp; i 3 about the same in crown area or a little larger than i 2 , about 1/2 or a little more 
its height, with larger central and smaller lateral accessory cusps, lying postero-externally to that 
tooth, separated from c 1 by a small diastema; pm 2 large, unreduced, its crown area similar to that 
or i 2 or a little less, slightly intruded but separating c 1 and pm 4 ; i l _ 3 not much imbricated, i 3 about 
twice the bulk of i 1 _ 2 ; pm 2 not usually much reduced, about 1/2-3/4 or more the crown area of 
pm 4 . 

Included taxa: aladdin, bactrianus, lacteus, nathusii (Fig. 2b); mediterraneus, (?) permixtus, 
pipistrellus (Fig. 2a). 

Among African Pipistrellus we have been unable to examine the baculum of permixtus (Aellen, 
1957) compared by its describer chiefly with nathusii. Its dentition, with bicuspid i 2 , the posterior 
cusp 2/3 the height of the anterior cusp, i 3 with lateral accessory cusps, its main cusp equal in height 
to the posterior cusp of i 2 , large, slightly intruded pm 2 which is about as big as i 3 , and unreduced 
pm 2 , its crown area about 3/4-4/5 the crown area of pm 4 suggests that it should be referred to the 
pipistrellus subgroup. Koopman (1975) referred it to the pipistrellus group. 

(a) (ii) javanicus (abramus) subgroup. Braincase slightly globular, elevated posteriorly; post- 
orbital region wide; supraorbital region distinctly broadened to produce abruptly incurving lateral 
margins to the anterior part of the postorbital area; rostrum broad, dorsally flattened, with no 
more than an indication of a median rostral depression; cranial profile almost straight from 
occiput to nares, slightly flattened over the occiput and a little depressed over the anterior part of 
the orbits; premaxillae not shortened; zygomata slender but not weak, lacking any jugal eminence; 
interdental palate only little longer than wide; palate strongly domed with broad anterior emargi- 
nation; maxillary toothrows more or less parallel, scarcely convergent anteriorly; short bony 
post-palate; shallow basial pits; i 2 well developed, bicuspid, posterior cusp sometimes small, 
usually about 3/4 height of anterior cusp; i 3 similar in size to i 2 or slightly larger, about as high as its 
posterior cusp, with larger central and smaller lateral accessory cusps, lying postero-externally to 
that tooth, separated from c 1 by a narrow diastema; pm 2 little reduced, equal to or rather less than 
i 3 in crown area, in recess between c 1 and pm 4 which approach but do not touch; ij_ 3 scarcely 
imbricated, i 3 as a rule similar in size to i 2 , both a little more massive than i^ pm 2 about 1/2-3/4 the 
size of pm 4 , very slightly intruded from toothrow. 

Included taxa: abramus (Fig. 3a), akokomuli, babu (Fig. 4a), bancanus, camortae (Fig. 15d), 
endoi (Fig. 3b), irretitus, javanicus (Fig. lOe), meyeni, peguensis (Fig. 15c), paterculus (Fig. 3c), 
pumiloides. 

Current treatments of Asian Pipistrellus usually include abramus in P. javanicus (tralatitius , 
Laurie & Hill, 1954) as a valid subspecies. There appear to be few conventional features that clearly 
separate javanicus from abramus but their bacula differ quite sharply in the high degree of vertical 
flexion of the shaft evident in the latter. This difference was used by Thomas (1928a) who examined 
Indo-Chinese Pipistrellus and differentiated abramus from raptor, javanicus (as tralatitius) and 
coromandra by virtue of the double curvature of its baculum, the others being straight. Van Peenen 
et al. (1969) recorded coromandra, javanicus and mimus from Vietnam but the baculum that they 
illustrate for javanicus is clearly that of abramus. This bacular difference suggests thatjavanicus and 
abramus should be considered specifically distinct even although there seem to be few cranial and 
dental characters to separate them. The braincase in javanicus is slightly more inflated than in 
abramus and its rostrum narrower, the palate is usually a little wider in relation to its length and is 
slightly more excavated and domed, while pm 2 is a little less reduced and less intruded, tending 
rather more to separate c 1 and pm 4 . Both occur in Vietnam (Thomas, 1928a; specimens listed 
below). It seems likely that bancanus and camortae, which has an unflexed baculum, are more 
closely related to javanicus than to abramus. 

Soota & Chaturvedi (1980) remarked that Thomas (191 5c) had pointed out that the baculum of 
abramus is doubly curved and that in paterculus it is straight, but they stated further that material of 



VESPERTILIONINE SYSTEMATICS 241 

paterculus in the collections of the Zoological Survey of India revealed that its baculum is doubly 
curved. However, specimens in the collections of the British Museum (Natural History) referred to 
paterculus (some the original material seen by Thomas) have relatively straight bacula when 
compared with the sinuous baculum of abramus. We have found this sinuous baculum to be 
characteristic of abramus, to which perhaps the specimens seen by Soota & Chaturvedi should be 
referred. 

The very elongate baculum of paterculus, with its strongly bifid tip, the 'horns' of which are 
deflected ventrally and extend to some extent to form a ring (Thomas, 191 5c) is reminiscent of the 
baculum of Scotoecus. A very long baculum is also found in endoi, but in this species the tip is 
less strongly bifid and the 'horns' are deflected dorsally. Both, however, are clearly referable to 
Pipistrellus on cranial and dental characters, Scotoecus being distinguished especially by a massive 
unicuspid i 2 , the loss of i 3 , a grooved c 1 , and usually by the absence of pm 2 . 

(a) (iii) coromandra subgroup. Small, with small, rounded braincase, elevated posteriorly and 
slightly so frontally; postorbital region wide; rostrum short, relatively narrow; no median rostral 
depression; cranial profile straight or nearly so from occiput to tip of rostrum; premaxillae excep- 
tionally short; zygomata slender, without jugal projection; interdental palate about as long or a 
little longer than wide; short bony post-palate: no basial depressions; i 2 usually bicuspid, posterior 
cusp sometimes very small or rarely absent, when present about 1/2 or a little more the height of the 
anterior cusp; i 3 equal or greater than i 2 in crown area, reaching to tip of its posterior cusp, with 
larger principal cusp and smaller lateral accessory cusps, lying postero-externally to the inner 
tooth; pm 2 not much reduced, nearly as great or as great in crown area as i 3 , with well developed, 
slightly inwardly directed pointed cusp, in recess between c 1 and pm 4 ; ^ _ 3 not much imbricated, i 3 
a little larger than i : _ 2 ; pm 2 about 1/2 crown area and height of pm 4 , slightly extruded. 

Included taxa: adamsi (Fig. lOc), afghanus, angulatus, collinus (Fig. 4b), coromandra (Fig. 7c), 
glaucillus, mimus (Fig. 7g), murrayi (Fig. 4c), nitidus, papuanus (Fig. 2c), ponceleti (Fig. 4d), 
portensis, principulus, sewelanus, sturdeei; possibly subulidens which may however represent 
javanicus; tenuis (Fig. 9d), tramatus (Fig. 7b), wattsi (Fig. lOg), westralis (Fig. lOd). 

(a) (iv) ceylonicus subgroup. Large, with rather short, broad braincase; wide postorbital region; 
some degree of supraorbital expansion; rostrum broad, rather long; weak, diffuse median rostral 
depression; cranial profile slightly convex, raised over the frontal region; premaxillae normal, not 
shortened; zygomata moderate, without jugal eminence or process, interdental palate longer than 
wide; maxillary toothrows parallel; short bony post-palate; slight basial pits; i 2 large and massive, 
bicuspid to almost unicuspid, with moderate to small posterior cusp about 2/3 height of anterior 
cusp; i 3 massive, as large or larger than i 2 , extending to or a little beyond posterior cusp of that 
tooth, with large principal cusp and smaller lateral accessory cusps, lying postero-laterally to i 2 , 
narrowly separated from c 1 ; pm 2 large, nearly as great or greater in crown area than i 3 , usually 
filling the recess between c 1 and pm 4 into which it is intruded, these almost in contact labially; i : _ 3 
slightly imbricated, i 3 a little larger than i 1 - 2 ' pm 2 almost as large in crown area as pm 4 , very 
slightly extruded from the toothrow. 

Included taxa: borneanus, ceylonicus (Fig. 7d), chrysothrix, indicus, (?) minahassae, raptor (Fig. 
3d), shanorum, subcanus. 

An account of minahassae is given by Tate (\942a) who referred it to a minahassae group of 
which it was the sole member. The skull of the holotype has never been described and Tate's 
remarks are based on a referred specimen in the American Museum of Natural History, New York 
(AMNH 102359). It has a short, high braincase with rudiments of a sagittal crest, prominent 
supraorbital tubercles and slender zygomata; i 2 is long, with well developed posterior cusp, c 1 
slender, lacking an accessory cusp, pm 2 only slightly intruded, its crown area greater than that of i 3 , 
and i^-j scarcely imbricated. These features suggest that if this specimen represents minahassae 
the taxon should be allocated to Pipistrellus (Pipistrellus) and provisionally we place it in the 
ceylonicus subgroup of the pipistrellus group, but clearly these decisions can only be speculative. 

(b) rueppellti group 

Baculum as in pipistrellus group; braincase high, broadened, rounded and globose; postorbital 



242 J. E. HILL & D. L. HARRISON 

region wide; supraorbital region slightly expanded; rostrum short; with shallow, ill-defined median 
depression; cranial profile almost straight, a little raised over frontal region, a little depressed over 
rostrum; premaxillae not shortened; zygomata slender, without jugal projection; interdental palate 
a little longer than wide; maxillary toothrows slightly convergent; short bony post-palate; no basial 
pits, instead a shallow depression; i 2 strongly bicuspid, posterior cusp about 3/4 height of anterior 
cusp; i 3 usually very small or minute, its crown area less than 1/2 that of i 2 , its tip sometimes barely 
rising above the cingulum of the inner tooth, on occasion (e.g. nanulus) larger, equal to or slightly 
exceeding i 2 in crown area, about 1 /2 or a little more the height of that tooth; i 3 lying sublaterally to 
i 2 , separated from c 1 by a wide diastema; pm 2 not usually greatly reduced, its crown area similar to 
that of i 2 , with strong cusp, separating c 1 and pm 4 , occasionally (crassulus) much reduced, similar 
in size to i 3 in its much reduced condition, or (crassulus, nanulus) recessed between these teeth; i t _ 3 
little imbricated, i 3 slightly the largest as a rule; pm 2 about 3/4 or more as large in crown area as 
pm 4 and about 3/4 its height, rarely (coxi, crassulus) more reduced, about 1/2 crown area and 
height of pm 4 . 

Included taxa: Probably coxi', crassulus (Fig. le),fuscipes, leucomelas, nanulus (Fig. lf),pulcher 
(Fig. lOa), rueppellii (Fig. \0b),senegalensis, vernayi. 

Vansonia Roberts, 1946 is available should further separation of the rueppellii group be thought 
justified: an earlier name, Alobus Peters, 1867 is preoccupied. 

(c) kuhlii group 

Baculum of moderate length with narrow cylindrical shaft and paired basal lobes as in pipistrellus 
and rueppellii groups but basal lobes strongly angled to line of shaft in vertical plane; braincase low 
but not flattened, rounded, only slightly elongate; postorbital region wide; supraorbital region not 
widened or swollen; rostrum long, unwidened, with very slight median flattening; cranial profile 
almost straight from occiput to nares, slightly raised over frontal region, slightly depressed over 
front of orbits; premaxillae slightly shortened; zygomata slender, weak, without jugal eminence; 
interdental palate longer than wide; maxillary toothrows almost parallel; short bony post-palate; 
small, narrow basial pits; i 2 usually unicuspid, at best only slightly bicuspid; i 3 small, its crown area 
1/2 or less that of i 2 , its tip extending only slightly beyond the cingulum of that tooth, to which it lies 
laterally or sublaterally, separated from c 1 by a moderate or narrow diastema; pm 2 small, similar in 
crown area to i 3 , intruded to lie in recess between c 1 and pm 4 , these more or less in contact; i t _ 3 
moderately imbricated, i 3 slightly the largest; pm 2 reduced, about 1/2 or less the crown area and 
height of pm 4 . 

Included taxa: Probably aero; deserti(aegyptius, Qumsiyeh, 1985) (Fig. 5c),fuscatus, ikwanius; 
probably inexspectatus; kuhlii (Fig. 5a), maderensis (Fig. 5b), marrensis, rusticus (Figs 5d, 6c). 

We have been unable to examine the baculum of inexspectatus (Aellen, 1959) but this taxon was 
placed in the kuhlii group by Koopman (1975) who also referred maderensis to the savii group. 
However, an example of maderensis in the collections of the British Museum (Natural History) has 
a baculum clearly of the kuhlii type. 

Romicia Gray, 1838 is available for the kuhlii group should this be thought worthy of further 
separation. 

Subgenus Pipistrellus ( Vespadelus) 

Baculum usually with long cylindrical or ventrally slightly fluted shaft, paired basal lobes and a 
blunt tip; shaft shorter and wider in sagittula; basal lobes sometimes flexed to line of shaft in 
vertical plane; braincase slightly broadened, flattened and elongated; postorbital region wide; 
supraorbital region slightly broadened; rostrum short but not greatly widened; shallow median 
rostral depression; cranial profile almost straight from occiput to nares, a little depressed over 
rostrum; premaxillae not shortened; zygomata slender, without jugal process; interdental palate a 
little longer than wide; maxillary toothrows slightly convergent anteriorly; short bony post-palate; 
no basial pits: i 2 bicuspid, posterior cusp almost as high as anterior cusp; i 3 much reduced, its 
crown area 1 /2 or less that of i 2 , its tip barely extending beyond the cingulum of that tooth, to which 
i 3 lies postero-laterally, separated from c 1 by a narrow diastema; pm 2 almost invariably absent, 



VESPERTILIONINE SYSTEMATICS 243 

when present a small spicule in recess between c 1 and pm 4 : i { _ 3 moderately imbricated, i 3 slightly 
the largest, pm 2 greatly reduced, in crown area about 1/2 or more usually less the crown area of 
pm 4 , and 1/2 its height. 

Included taxa: caurinus (Fig. lib), douglasorum (Fig. 1 ld),pumilus (Figs 1 la, 12k), regulus (Fig. 
1 le), sagittula (Fig. 1 1 0, vulturnus (Fig. 1 Ic). 

Formerly referred to Eptesicus, the transfer of these taxa to Pipistrellus was first suggested by 
Heller & Volleth (1984), purely on bacular grounds. The bacular, cranial and dental features of this 
group suggest that it represents P. (Pipistrellus) in Australia, the few members of this subgenus 
(adamsi, westralis and perhaps javanicus) that also occur there being possibly slightly less differen- 
tiated by virtue of their relatively slightly less shortened skulls and their retention of pm 2 . The 
pipistrellus group of the subgenus extends widely through the islands of Indo- Australia to New 
Guinea, the Solomon Islands and Australia, chiefly as the coromandra subgroup, to which adamsi 
and westralis belong. Thejavanicus subgroup reaches at least to Java and Sulawesi and may extend 
to Australia (Hill, 1983) but the Australian record ofjavanicus is based on two old examples and 
has never been confirmed. Possibly the slightly differentiated pumilus and it allies result from a 
further perhaps earlier invasion of Australia. Bacular differences in this subgenus (Figs 11, 12k) 
suggest that it may consist of two groups: it has been possible to examine only pumilus. 

There has been hitherto a wide geographical hiatus in the Indo-Australian distribution of 
Eptesicus as formerly understood. Beyond these Australian forms, no other taxon attributed to 
this nominal genus has been reported further east in Indo-Australia than southern Thailand, other 
than an unconfirmed record from Sarawak of Eptesicus sp. (Pirlot, 1968) which provided no 
details. 

Subgenus Pipistrellus (Perimyotis) 

Baculum very small, Y-shaped, with paired basal lobes and very short shaft; braincase slightly 
elongate, rounded, almost globose; postorbital region wide; supraorbital region slightly broad- 
ened; rostrum long, elevated, slightly widened; shallow median frontal depression; a very slight 
lateral depression on each side just anterior to the orbital rim; cranial profile sinuous, raised 
over frontal region, a little depressed over front of orbits; premaxillae not shortened; zygomata 
moderate, a slight jugal eminence; interdental palate longer than wide with wide anterior palatal 
emargination; maxillary toothrows convergent anteriorly; very short bony post-palate; slight 
basial depressions; i 2 bicuspid with well developed posterior cusp about 3/4 height of anterior cusp; 
i 3 massive, its crown area exceeding that of i 2 , in height reaching or exceeding the height of anterior 
cusp of that tooth, with larger principal cusp and smaller lateral accessory cusps, lying postero- 
externally to inner tooth, separated from c 1 by a wide diastema; pm 2 large, its crown area equal to 
that of i 3 , in toothrow, sometimes separated from pm 4 by a slight diastema; i 2 " 3 and pm 2 almost 
identical to those of P. nathusii; i^_ 3 not imbricated, i 3 only slightly bulkier than i : _ 2 ; pm 2 not 
greatly reduced or compressed in toothrow, its crown area about 1/2 or more that of pm 4 , about 
1/2-3/4 its height; tragus myotine, about 1/2 height of ear, tapering to blunt point. 

Included taxon: subflavus (Fig. 2d). 

Menu (1984) proposed the genus Perimyotis for P. subflavus, chiefly on account of the features of 
the canine and post-canine dentition in which he believed this species to approach Myotis. How- 
ever, Hill & Topal (1973) in discussing Myotis rosseti and M. ridleyi which also combine the tragal 
features of Myotis with the Pipistrellus dentition (pm| absent) noted that in Myotis i 2 is short and 
broad, its posterior cusp wider basally than the anterior cusp, while in Pipistrellus this tooth is 
linear, often narrower posteriorly than anteriorly. Also, in Myotis the principal cusp of i 3 is equal 
to or exceeds that of i 2 in height and the tooth is often hooked to produce a caniniform apperance 
while in Pipistrellus it is lower and is not hooked. In Myotis i 3 is usually much larger than i l _ 2 but 
in Pipistrellus there is as a rule no such great distinction in size. The incisive dentition of subflavus 
corresponds closely with that of Pipistrellus. 

The baculum of subflavus is of a type not found in Myotis. Menu (1984) stated on the basis of 
published figures that the baculum approached that of certain Myotis and more particularly that of 
Plecotus auritus. We find no significant resemblance to the morphologically rather stable, saddle- 



244 J. E. HILL & D. L. HARRISON 

like baculum ofMyotis and although there are some similarities with the bacula ofPlecotus auritus 
(Fig. 19g) and P. teneriffae (Fig. 14d), that of P. austriacus (Fig. 19h) is nearer in structure to the 
myotine baculum. The bacular type found in subflavus occurs in a similar form in Pipistrellus 
circumdatus (Fig. 2e), P. societatis (Fig. 9c) and P. cuprosus (Fig. 9h). There are considerable 
differences, however, between subflavus and circumdatus and its allies, not least in the degree of 
reduction of pm 2 , this tooth in these three species being very small or absent. 

The unshortened rostrum and the dental features of subflavus suggest that it is nearest to P. 
(Pipistrellus), which it appears to represent in North America. We find hesperus, the other North 
American species of Pipistrellus, to belong on bacular and dental grounds to P. (Hypsugo). Thus 
we do not support Menu's view (p. 410, footnote) that Pipistrellus is limited to the Old World and 
that the lines leading to this genus did not enter the North American continent. The marked 
differences between subflavus and hesperus indicate two quite different pipistrelline groups, as 
Hamilton (1949) remarked in relation to their bacula, but bacular and dental evidence suggests 
alliance to established Old World groupings, the baculum of subflavus being perhaps a reduced 
form of the shafted structure found in P. (Pipistrellus) , that of hesperus a modification of the type 
found in P. (Hypsugo) . 



Subgenus Pipistrellus (Hypsugo) 

Baculum usually short, stout, sometimes with expanded base and tip; base rarely bilobed, some- 
times dorsally elevated; shaft generally flattened dorso-ventrally, sometimes wide, its underside 
transversely concave or fluted; tip ventrally hollowed as an extension of ventral fluting of shaft, 
when expanded anteriorly sub-square or slightly rounded, its anterior edge sometimes irregularly 
serrated, tip sometimes downwardly directed, its lateral margins on occasion forming two broadly 
based, ventrally directed projections; pm 2 generally much reduced, small, minute, or rarely absent. 
Wallin ( 1 969) considered Hypsugo a valid subgenus within Pipistrellus but included only P. savii: 
Horacek & Hanak (1985#, b) added cadornae and pulveratus and suggested the elevation of 
Hypsugo to generic rank, subsequently (1985-1986) widening its possible content and considering 
it generically distinct. 

(a) savii group 

Postorbital region, supraorbital region and rostrum not greatly widened; supraorbital tubercles if 

present small and undeveloped. 

(a) (i) pulveratus subgroup. Braincase elongate, inflated; postorbital region wide, supraorbital 
area not broadened; rostrum long, not widened; shallow frontal depression; no median rostral 
depression; broad, shallow lateral depressions above anterior part of orbit; cranial profile 
somewhat sinuous, depressed over front of orbits; premaxillae not shortened; zygomata robust, 
with very slight jugal eminence; interdental palate longer than wide; maxillary toothrows almost 
parallel; moderate bony post-palate; no basial pits; i 2 bicuspid, posterior cusp about 3/4 height of 
anterior cusp; i 3 large, wide, its crown area equal to or slightly exceeding that of i 2 , reaching to tip 
of the posterior cusp of that tooth, with moderate lateral accessory cusps, lying postero-externally 
to the inner tooth, separated from c 1 by a moderate diastema; pm 2 about equal or nearly equal to 
i 3 in crown area, in recess between c 1 and pm 4 which are closely approximated; ^.3 slightly 
imbricated, i 3 a little the largest; pm 2 a little less than 1/2 the crown area of pm 4 , 1/2-3/4 its height. 
Included taxon: pulveratus (Fig. 8c). 

(a) (ii) nanus subgroup. Braincase elevated, slightly inflated, more or less globose but a little 
elongated; postorbital region wide; supraorbital area slightly widened with small supraorbital 
swellings; rostrum not especially shortened or broadened; shallow median rostral depression; 
slight lateral depressions just anterior to supraorbital region; cranial profile sinuous, strongly 
depressed and concave over rostrum; premaxillae not shortened; zygomata slender, lacking jugal 
projection; interdental palate longer than wide; maxillary toothrows slightly convergent; short 
bony post-palate; no basial pits; i 2 unicuspid or with small posterior cusp extending to about 3/4 of 



VESPERTILIONINE SYSTEMATICS 245 

its height; i 3 wide, its crown area slightly exceeding that of i 2 , about 1/2-3/4 the height of that tooth, 
extending almost to the tip of its posterior cusp, with slight lateral cusps, lying postero-externally 
to the inner tooth, separated from c 1 by a wide diastema; pm 2 about 1/2-2/3 or a little more the 
crown area of i 3 , intruded into recess between c 1 and pm 4 , these sometimes in contact or nearly so; 
i : _ 3 not or only very slightly imbricated, i 3 slightly the largest; pm 2 about 1/2 the crown area and 
height of pm 4 . 

Included taxa: arabicus (Fig. 7a), culex, Helios (Fig. 6d); probably musciculus; nanus (Fig. 6b), 
stampflii. 

Current listings (i.e. Hayman & Hill, 1971; Koopman, 1975) unite Helios with P. nanus as a 
synonym or possibly as a valid subspecies. However, the bacular features of this pale form suggest 
that it may represent a species distinct from nanus with which it may be sympatric in northern and 
eastern Kenya and in the Sudan. No baculum has been available for musciculus, which was placed 
in a Hesperus group by Koopman (loc. cit.). Although its incisive and premolar dentition agrees 
with the nanus subgroup its placement here remains speculative. 

(a) (in) savii subgroup. Braincase rather low and flat, elongate rather than globose; postorbital 
region not especially widened; supraorbital region unwidened or only slightly widened; rostrum of 
moderate length; a shallow median rostral depression; usually slight lateral rostral depressions just 
anterior to supraorbital and anterior orbital rim; cranial profile straight or slightly concave; 
premaxillae not shortened; zygomata robust, often with slight jugal process; interdental palate a 
little longer than wide; maxillary toothrows more or less parallel; short bony post-palate; shallow 
or no basial pits; i 2 unicuspid or with posterior cusp, when present about 3/4 height of anterior 
cusp; i 3 similar to or exceeding i 2 in crown area, about 1/2-3/4 the height of i 2 , with strong 
central cusp flanked by smaller lateral accessory cusps, lying postero-externally or more laterally 
(anchietae) to the inner tooth, separated from c 1 by a strong diastema; pm 2 much reduced, minute 
or absent, when present crown area less than 1/2 that of i 3 , in recess between c 1 and pm 4 , these as a 
rule in contact; \^ _ 3 slightly or more strongly imbricated, similar in size or i 3 slightly the bulkiest; 
pm 2 reduced, about 1/2 or less in crown area than pm 4 and about 2/3 its height. 

Included taxa: anchietae (Fig. 6e); probably ariel', probably austenianus; bodenheimeri (Fig. 9f), 
caucasicus, darwini, maurus, savii (Fig. 6a). 

We have been unable to examine the baculum of ariel. The baculum of a small Pipistrellus from 
the Naturhistorisches Museum, Wien (from Sayala, Upper Egypt) tentatively identified as ariel is 
illustrated by Gaisler et al. (1972) but is evidently of the kuhlii type. Qumsiyeh (1985) employs the 
description of this baculum in his account of ariel. However, Dr K. Bauer informs us (in litt.) that 
the specimen (NHW 10351) of which the baculum is figured by Gaisler et al. (loc. cit.) is not 
referable to ariel but is instead a small deserti, an identification clearly supported by its bacular 
structure. Moreover, Dr Bauer has loaned three similarly small specimens, one male, the others 
female (NHW 27501-3) (length of forearm 29-2, 28-9, 28-2; condylobasal length 11-0, 10-5. 10-9; 
c-m 3 4-0, 3-8, 3-9) apparently from Upper Egypt, that also represent deserti: a baculum from this 
sample is again exactly of the kuhlii type. The cranial (narrow braincase, unexpanded rostrum, 
short broad narial and anterior palatal emarginations, narrow basioccipital) and dental (long i 3 , 
minute pm 2 ) features of ariel clearly indicate that it belongs with savii, to which group Koopman 
(1975) referred it. 

\syntype of Eptesicusbicolor(Bocage, 1889)(BM(NH) 89.5. 1.3) (Fig. 9e) proves to be identical 
cranially, dentally and in bacular morphology with Pipistrellus anchietae (Seabra, 1900) (vide 
infra, p. 249). However, the point needs confirmation or otherwise by examination of the other 
syntype in the Museu Nacional de Lisboa. It should be noted that bicolor is the prior name 
(HonackietaL, 1982). 

The relationship between the pulveratus , nanus and savii subgroups is illustrated by arabicus and 
bodenheimeri, the bacula of which are compared directly by Harrison (1982). The baculum of 
arabicus (Fig. 7a) approaches that of anchietae (Fig. 6e) yet cranially and dentally this species is 
nearer to nanus (Fig. 6b), while that of bodenheimeri (Fig. 9f) is like the baculum of pulveratus (Fig. 
8c) but cranially and dentally the species is close to savii (Fig. 6a). These combinations of features 
link the three subgroups. 



246 J. E. HILL & D. L. HARRISON 

(a) (iv) Hesperus subgroup. Baculum a fluted structure, much like that ofpulveratus or bodenhei- 
meri. Braincase low but broad, elongated; postorbital region wide; supraorbital area slightly 
widened; rostrum short, not greatly broadened; a shallow median frontal depression; slight lateral 
rostral depressions just above anteorbital foramina; cranial profile almost straight, slightly 
depressed above anterior root of zygomata; premaxillae not shortened; zygomata slender, a little 
widened anteriorly, lacking any jugal eminence; interdental palate about as wide as long; maxillary 
toothrows convergent; short bony post-palate; no basial pits; cochlear bullae inflated with narrow 
basioccipital; i 2 unicuspid; i 3 slightly greater in crown than i 2 but about 1/2 its height, with little 
trace of lateral accessory cusps, lying postero-externally, separated from c 1 by moderate to small 
diastema; pm 2 small to minute, at best about 1/2 or less in crown area than i 3 , in recess between c 1 
and pm 4 which are closely approximated; ij_ 3 scarcely or not imbricated, similar in size; pm 2 
reduced, about 1/2 the crown area of pm 4 , a little less than 1/2 its height. 

Included taxon: Hesperus (Fig. 8d). 

Horacek & Hanak (1985a, b, 1985-1986) have indicated that they intend to propose generic 
status for Hesperus and indeed have suggested that it be referred to Parastrellus which they offer as a 
new name. It is however a nomen nudum in these publications. There seem good grounds for 
considering Hesperus the North American representative of P. (Hypsugo) to which its bacular, 
cranial and dental features ally it. Like bodenheimeri (Fig. 9f) its baculum approaches that of 
pulveratus (Fig. 8c) but cranially and dentally it is nearer to savii&nd its immediate allies. Koopman 
(1975) referred Hesperus to a Hesperus group in which he also included the African musciculus, here 
provisionally allocated to the nanus subgroup. 

(a) (v) eisentrauti subgroup. Braincase broad, elevated and globular; inflated frontally; post- 
orbital region wide; supraorbital region broadened, with small supraorbital tubercles; rostrum 
short, deep, wide and massive; slight median rostral depression; cranial profile straight or slightly 
convex; premaxillae not shortened; zygomata strong, lacking any jugal projection; interdental 
palate very slightly longer than wide; maxillary toothrows almost parallel; short bony post-palate; 
slight basial pits usually present; i 2 long, narrow, bicuspid, posterior cups about 3/4 height of 
anterior cusp; i 3 short, wide, similar to or slightly greater in crown area than i 2 , about 1/2 or a little 
more its height, with larger central cusp and smaller lateral accessory cusps, lying laterally and 
slightly posteriorly to the inner tooth, separated from c 1 by a moderate diastema; pm 2 small, about 
the same in crown area as i 3 , sandwiched into recess between c 1 and pm 4 , these almost in contact; 
i t _ 3 slightly imbricated, i 2 _ 3 similar in size, both larger than i t ; pm 2 about 1/2 crown area and 
height of pm 4 . 

Included taxon: eisentrauti (Fig. 9g). 

Koopman (1975) places eisentrauti in a rueppellii group, no doubt on account of its elevated, 
inflated braincase and its bicuspid i 2 , but its bacular features do not associate it with this species 
and its immediate allies. Its baculum is very similar to that of imbricatus (Fig. 9a) or macrotis 
(Fig. 9b). 

(a) (vi) imbricatus subgroup. Braincase inflated, globular, raised posteriorly; postorbital region 
Wide; supraorbital area slightly widened with very small supraorbital tubercles; rostrum short, 
not especially broadened; no median rostral depression; cranial profile almost straight; slightly 
concave above supraorbital region; premaxillae not shortened; zygomata moderate to strong, 
sometimes with a trace of a jugal eminence; interdental palate about as wide as long, not domed; 
maxillary toothrows almost parallel; very short bony post-palate; well developed basial pits; i 2 
bicuspid, posterior cusp about 3/4 height of anterior cusp; i 3 similar in crown area to i 2 , about 1/2 
its height, with larger central cusp and smaller lateral accessory cusps, lying laterally to the inner 
tooth, separated from c 1 by a narrow diastema; pm 2 greatly reduced, 1/4 or less the crown area of 
i 3 , in recess between c 1 and pm 4 , these in contact; i l _ 3 scarcely imbricated, i 2 _ 3 of similar size, a 
little larger than i t ; pm 2 about 1/2 the crown area and height of pm 4 . 

Included taxa: curtatus, imbricatus (Fig. 9a), macrotis (Fig. 9b), vordermanni. 

(a) (vii) lophurus subgroup. Braincase inflated, rounded, slightly elongate, raised posteriorly; 
postorbital region wide; supraorbital area little widened; at best only a trace of supraorbital 



VESPERTILIONINE SYSTEMATICS 247 

tubercles; rostrum moderate in length, longer than in imbricatus subgroup, not broadened; no 
median rostral depression; cranial profile almost straight, slightly depressed or concave above 
supraorbital region; zygomata strong with distinct jugal eminence; interdental palate a little longer 
than wide; maxillary toothrows slightly convergent; moderate bony post-palate; deep basial pits; 
incisor and premolar dentition closely similar to that of imbricatus subgroup but i 3 lying more 
postero-laterally to i 2 , and pm 2 sometimes (lophurus) slightly larger, about 1/2 crown are of i 3 . 

Included taxa: cadornae, kitcheneri (Fig. 8e), lophurus (Fig. 80- 

The baculum of kitcheneri is unusual in the presence distally of two anterior dorso-lateral, 
posteriorly directed processes, with ventrally a more or less tapered median gutter. As in lophurus, 
the tip is directed slightly ventrally. 

(b) stenopterus group 

Braincase large, rounded and globular; postorbital region very wide; supraorbital region much 
widened to include well developed supraorbital tubercles; rostrum short, wide; shallow median 
rostral depression anterior to frontal region; cranial profile slightly convex, elevated over frontal 
area; premaxillae not shortened; zygomata rather weak, lacking jugal process but usually with 
small descending process external to m 3 ; palate short and broad, the interdental palate as wide as 
long; maxillary toothrows parallel or nearly so; short bony post-palate; shallow basial pits; i 2 
small, bicuspid, posterior cusp 1/2-3/4 height of anterior cusp; i 3 a little smaller in crown area than 
i 2 , its tip reaching almost to tip of the posterior cusp of that tooth, with large central cusp and 
smaller lateral accessory cusps, lying postero-laterally to the inner tooth, only narrowly separated 
from c 1 or almost in contact with it; c 1 with distinct, well defined posterior accessory cusp; pm 2 
small or minute, about equal in crown area or a little larger than i 3 (stenopterus) or about 1/3-1/4 
the crown area of this tooth (joffrei, anthonyi), in recess between c 1 and pm 4 , which touch; i t _ 3 not 
much imbricated, all of similar size; crown area of pm 2 slightly exceeding that of pm 4 , pm 2 similar 
in height to the second tooth (stenopterus), or crown area of pm 2 about 1/2 that of pm 4 , pm 2 almost 
as high as that tooth (joffrei, anthonyi). 

Included taxa: anthonyi, joffrei, stenopterus (Fig. 7h). 

The baculum of stenopterus is unusual, although of the savii type: it has a narrow lobed base, 
hollowed shaft, and expanded tip the lateral margins of which project ventrally as two broadly 
based 'horns'. The stenopterus group as here understood is the joffrei group of Tate ( 1 942a) and (in 
part) of Koopman (1973). Both joffrei and stenopterus have been referred in the past to Nyctalus 
but as mentioned above (p. 234) the baculum of stenopterus has no resemblance to the long-shafted 
baculum of that genus (Fig. 100- Tate (19420) referred both to Pipistrellus with the comment that 
the group approached Oriental members of the savii group, and might at a later time be accorded 
generic rank. 

Subgenus Pipistrellus (Falsistrellus) 

Baculum a broad, proximally widened and ventrally deeply fluted structure with no distal expan- 
sion; braincase elongate; postorbital region wide; supraorbital area not expanded; rostrum long, 
not broadened; zygomata moderate to strong; and palate rather narrow, the interdental palate 
longer than wide. 

Pipistrellus (Falsistrellus) appears to be related to P. (Hypsugo) of which it may be the eastern 
representative. It is approached in bacular morphology by some of the latter subgenus such as 
imbricatus (Fig. 9a), macrotis (Fig. 9b), kitcheneri (Fig. 8e) and lophurus (Fig. 8f), and indeed the 
baculum in P. (Falsistrellus) appears to be an extreme variant of the broad, ventrally fluted 
structure of many of P. (Hypsugo). 

Kitchener et al. (1986) raised Falsistrellus to generic rank but did not include affinis andpetersi, 
confining their comparisons to the Australasian Pipistrellus (i.e. adamsi, angulatus, collinus, 
papuanus, wattsi and westralis) here referred to the coromandra subgroup of P. (Pipistrellus). These 
authors drew attention to its larger size; to its small i 3 which is anteriorly displaced and swivelled or 
rotated outwards to lie alongside i 2 , its concavity facing outwards (a feature which may have 
influenced Iredale & Troughton, 1934 in placing it in Glischropus); and to its combination of 



248 J. E. HILL & D. L. HARRISON 

unicuspid i 2 , tiny pm 2 and pronounced occipital crest, which asTate (19420) noted gives the rear of 
the skull a 'helmeted' appearance. Excepting the large size and the presence of a strong occipital 
crest, these features occur elsewhere in the various groups of Pipistrellus: the extent of the occipital 
crest may be a function of the large size of the skull. 

(a) affinis group 

Braincase rather narrow, mastoid width markedly less than zygomatic width; postorbital region 
wide; slight, rather poorly developed supraorbital ridges; very shallow median rostral depression 
just anterior to frontal region; dorso-lateral margin of rostrum more or less straight from post- 
orbital constriction to anterior edge of orbit; cranial profile almost straight, slightly elevated 
frontally, depressed over supraorbital region; premaxillae not shortened; zygomata moderate to 
robust with jugal eminence; maxillary toothrows slightly convergent; moderate bony post-palate; 
no basial pits; i 2 strongly bicuspid, posterior cusp 3/4 or more the height of the anterior cusp; i 3 
larger in crown area than i 2 , its height about equal to the height of the posterior cusp of that tooth, 
with strong central cusp and smaller lateral accessory cusps, lying postero-laterally to the inner 
tooth, separated from c 1 by a moderate diastema; pm 2 almost as great in crown area as i 3 (affinis) or 
about 3/4 its crown area (petersi), in recess between c 1 and pm 4 ; i^ _ 3 moderately imbricated, i 3 the 
largest; pm 2 about 1/2 the crown area of pm 4 and about 2/3-3/4 its height. 

Included taxa: affinis (Fig. 8a), (?) mordax, petersi (Fig. 8b). 

It has not been possible to examine mordax. Indian records of this species appear to be based on 
specimens in the collections of the British Museum (Natural History) tentatively labelled as such. 
These, however, agree closely with the description of P. affinis by Dobson (1871) and with the 
account of a specimen referred to this species from Likiang, Yunnan by Tate (19420). If correctly 
allocated, mordax (Peters, 1867) is the earliest name in the group. 

(b) tasmaniensis group 

Large and distinctive; braincase high, with well developed sagittal crest; postorbital region wide 
and strong; no median rostral depression; cranial profile straight; premaxillae slightly shortened; 
zygomata strong with slight jugal process and small inferior process; maxillary toothrows nearly 
parallel; short bony post-palate; slight basial depressions; i 2 large, unicuspid; i 3 small, its crown 
area about 1 /4 that of i 2 , barely extending above the cingulum of that tooth to which it lies laterally, 
its hollowed face outwardly directed, separated from c 1 by a moderate diastema: pm 2 very small, 
about 1 /3 the crown area of i 3 , in recess between c 1 and pm 4 , which touch; i l _ 3 much imbricated, i 3 
twice the bulk of i l - 2 \ pm 2 much reduced, about 1/4 the crown area of pm 4 and about 1/2 its 
height. 

Included taxa: mackenziei (Fig. lOh), tasmaniensis (Fig. 8g) 

Subgenus Pipistrellus (Neoromicia) 

Baculum with distinct paired basal lobes, slender cylindrical shaft and variously expanded tip; 
braincase broad, sometimes slightly elongate, rather flattened; postorbital region wide; supra- 
orbital area unwidened or only slightly broadened; rostrum moderate or slightly lengthened; 
cranial profile straight; premaxillae shortened; zygomata moderate, no jugal process; maxillary 
toothrows only slightly convergent; short bony post-palate; no basial pits; i 2 unicuspid or with 
small posterior cusp extending for about 3/4 its height; i 3 smaller than i 2 , its crown area 3/4-1/2 or 
less that of the inner tooth, about 1/2 its height, with usually a larger central cusp and slight lateral 
accessory cusps, the inner cusp as a rule very small, the tooth anteriorly displaced to lie alongside or 
almost alongside i 2 , separated from c 1 by a moderate to wide diastema; pm 2 almost invariably 
absent, when present very small, in recess between c 1 and pm 4 , i l _ 3 slightly to moderately imbri- 
cated, of similar size of with i 2 _ 3 a little the larger; pm 2 reduced, its crown area 1/2 or less that of 
pm 4 and its height 1/2-3/4 of the height of that tooth. 

This subgenus is wholly African and hitherto its members have been referred to Eptesicus, 
although there is karyological evidence (vide infra) suggesting that one at least should be moved to 
Pipistrellus. It incorporates the Eptesicus capensis and E. tenuipinnis groups of Koopman (1975). 



VESPERTILIONINE SYSTEMATICS 249 

These can be recognised readily by the structure of the baculum, capensis and its allies (Fig. 12a-d, 
f-i) having the distal part of the baculum spatulate and ventrally deflected, tenuipinnis and its 
associates (Fig. 12e, j) having the baculum modified distally into a lobed, almost vertical plate-like 
structure. 

Published karyological data refers only to capensis, although studies of other members of the 
subgenus are in progress (Rautenbach & Schlitter, 1985a, b). Peterson & Nagorsen (1975) found 
that capensis has a diploid number of 32 and a fundamental number of 50: Williams & Mares 
(1978) discussed the possible composite nature of Eptesicus as suggested by Koopman (1975) and 
pointed out that the species fitted karyologically within the variation exhibited by Pipistrellus. This 
genus has a diploid number varying from 26 to 44, and fundamental numbers from 44 to 60, these 
findings apparently supporting Koopman's observations. These authors remarked, however, that 
the karyotype of capensis is more similar to that of Pipistrellus nanus (2N = 36, FN = 50) than to P. 
kuhlii (2N = 44, FN = 50), Koopman having thought capensis nearer to the kuhlii group than to the 
pipistrellus group in which he placed nanus. Williams & Mares (loc. cit.) also found, in contrast, 
that small Eptesicus from the New World (diminutus,furinalis) have the typical 'eptesicoid' karyo- 
type (2N = 50, FN = 48-50), and added that the karyotypic differences between Eptesicus (sensu 
stricto) and Pipistrellus might prove more useful for separating these genera than other structural 
features. 

Our study of the bacula of African 'Eptesicus ' confirms these observations and indicates the 
isolation of capensis, tenuipinnis and their relatives from Eptesicus sensu stricto (vide infra): Heller 
& Volleth (1984) also transferred capensis to Pipistrellus, entirely on account of its published 
karyology. It is interesting to note also that the baculum of P. nanus indicates that this species 
should be referred to P. (Hypsugo) rather than to P. (Pipistrellus) where Koopman (1975) effec- 
tively allocated it. The bacular morphology of capensis, tenuipinnis and their allies suggests 
strongly that these former groupings of Eptesicus are most closely allied to P. (Hypsugo) as the 
karyological similarity of capensis to P. nanus indicates. The anterior upper premolar (pm 2 ) is very 
small, vestigial or absent in P. savii and is very small in most other members of P. (Hypsugo): very 
rarely it is present in capensis (Wallin, 1969; Hill & Topal, 1973). On the same point, we have been 
able to examine a specimen (MJS 2846) from Somalia, in the Carnegie Museum of Natural 
History, which has a small pm 2 on both sides of the jaw, leading to its erstwhile identification as 
Pipistrellus deserti. The baculum, however, is characteristically that of 'Eptesicus' somalicus, which 
in fact the specimen represents. 

Koopman (1975) suggested that Vesperus bicolor Bocage, 1889 ( = Eptesicus bicolor) and 
Pipistrellus anchietae (Seabra, 1900), both from Angola, may be conspecific, having examined 
syntypes of both at the British Museum (Natural History). This author thought that bicolor might 
be a form of 'Eptesicus ' tenuipinnis as Hayman & Hill (1971) suggested, or that it might be based on 
a specimen of Pipistrellus anchietae with missing anterior upper premolars. Bocage (loc. cit.) says 
'pas de trace de la premiere premolaire a la machoire superieure'. Further study of the syntype 
(BM(NH) 89.5. 1 .3) in London shows it to have a small pm 2 in a recess between c 1 and pm 4 on each 
side: cranially it agrees exactly with the syntype of anchietae (9 BM(NH) 6. 1 .3. 1) and its baculum is 
exactly as in that species. Curiously, Bocage states that both original specimens of bicolor are 
female. The specimen in London is quite clearly listed as a 'Co-type' by Thomas in the relevant 
accession register. 

(a) capensis group 

Tip of baculum flattened, deflected ventrally, sometimes a small sub-apical dorsal projection; 
braincase flattened, slightly elongate; rostrum not especially broadened; palate long, narrow, 
interdental palate longer than wide; i 3 1/2 or less the crown area of i 2 . 

Included taxa: capensis (Fig. 12b, g); probably brunneus, garambae, grandidieri; guineensis 
(Fig. 12c), matroka (Fig. 12a), melckorum (Fig. 12f); minutus (?) (Fig. 12i); probably rectitragus; 
somalicus (Fig. 12h); probably vansoni; zuluensis (Fig. 12d). 

The baculum of brunneus sensu stricto has not been examined. That (Fig. 14b) of a Nigerian 
specimen (BM(NH) 48.702) collected by I.T. Sanderson and hitherto referred to this species is 
very similar to that of rendalli (Fig. 12e), with which this example agrees in cranial and ventral 



250 J. E. HILL & D. L. HARRISON 

characters. The series whence this specimen comes is discussed by Koopman ( 1 965) and Hayman & 
Hill (1971). 

Our study of bacula in this group shows clearly that mat r oka belongs with capensis: we have been 
unable to examine the baculum of humbloti. We find too that capensis and somalicus can be 
separated by bacular features: the baculum of capensis has distally a downwardly directed, plate- 
like expansion, while in somalicus the distal part of the baculum is more spatulate, depressed just 
below the line of the shaft. Moreover, zuluensis is very clearly of the somalicus type, and the two 
appear to be very closely related, as Koopman (1975) suggested. Bacular morphology also con- 
firms the observation by this author that melckorum is like a giant capensis: Rautenbach & Schlitter 
(19850, b) suggested that these are synonymous. 

(b) tenuipinnis group 

Tip of baculum expanded into an almost vertical, lobed, plate-like structure; braincase similar to 
capensis group, but broader and less elongate; rostrum slightly widened; palate short and broad, 
interdental palate about as long as wide; i 3 about 1/2-3/4 the crown area of i 2 . 

Included taxa: Probably angolensis,faradjius,flavescens,phasma; rendalli (Fig. 12e), tenuipinnis 
(Fig. 12j). 

Pipistrellus (Arielulus) subgen. nov. 

TYPE SPECIES: Vespertilio circumdatus Temminck, 1840. Java. 
REFERRED SPECIES: Pipistrellus societatis Hill, 1 972; Pipistrellus cuprosus Hill & Francis, 1 984. 

DISTRIBUTION: Burma to Java (circumdatus, Fig. 2e); Malaya (societatis, Fig. 9c); Borneo 
(cuprosus, Fig. 9h). 

DIAGNOSIS: Differs from most other subgenera of Pipistrellus in very small, Y-shaped baculum 
which has paired basal lobes and a short shaft; baculum similar to that of P. (Perimyotis) but 
differing from this subgenus in greatly reduced i 3 and pm 2 , the former displaced anteriorly to lie 
alongside i 2 , the latter sometimes absent. 

DESCRIPTION: Size small to medium (length of forearm 34-7-43-6); muzzle short, broad and blunt; 
ears large, rounded, with blunt tip, anterior margin with prominent, posteriorly directed basal 
lobe, posterior margin with wide quadrate lobe at insertion just behind angle of mouth; tragal 
margin concave anteriorly, rising to anteriorly directed point, upper margin of tragus nearly 
horizontal, posterior margin strongly convex; ears and upper margin of tragus edged to a greater or 
lesser extent with dull white or yellowish white; dorsal pelage black or blackish brown, the hairs 
tipped with yellowish, orange, russet, copper or bronze. 

Braincase high, inflated, globose; postorbital region wide; supraorbital area broadened, with 
small supraorbital projections or tubercles; rostrum short, widened, sometimes a shallow median 
rostral depression; cranial profile almost straight, elevated frontally, slightly depressed behind and 
above supraorbital region; premaxillae not shortened; zygomata strong, no jugal eminence; inter- 
dental palate longer than wide; short to moderate bony post-palate; shallow to moderate basial 
pits; i 2 almost unicuspid, posterior cusp if present insignificant; i 3 very small, about 1/4 the crown 
area of i 2 , 1/3 or less its height, lying almost alongside this tooth, separated from c 1 by a narrow 
to moderate diastema; pm 2 very small or absent, when present similar in size to i 3 , recessed into 
angle between c 1 and pm 4 which are in contact; i t _ 3 considerably imbricated, \^_ 2 tricuspid or 
incipiently quadricuspid, i 2 _ 3 bulkier, larger than i 1 , similar in size to each other, their cusps 
indistinct; pm 2 1/2-1/4 size of pm 4 , compressed in toothrow. 

ETYMOLOGY: The new subgeneric name is a diminutive of Ariel, a little sprite. 

REMARKS: Heller & Volleth (1984) transfer circumdatus and societatis to 'Eptesicus' on karyo- 
logical, bacular and dental grounds. However, the baculum in these species does not resemble 
closely any of those found either in Eptesicus sensu lato or Eptesicus sensu stricto. Possibly the 
unusual baculum in P. (Arielulus}, similar to that of P. (Perimyotis), is a reduced form of the P. 



VESPERTILIONINE SYSTEM ATICS 2 5 1 

(Pipistrellus) type, but the species allocated to P. (Arielulus) differ widely cranially and dentally 
from P. subflavus, the sole species referred to P. (Perimyotis). 

Genus Eptesicus Rafinesque, 1 820 

Eptesicus Rafinesque, 1820: 2. Eptesicus melanops Rafinesque = Vespertilio fuscus Palisot de Beauvois. 

Cnephaeus Kaup, 1820: 103. Vespertilio serotinus Schreber. 

Noctula Bonaparte, 1837: fasc. xxi. Noctula serotina Bonaparte. 

Cateorus Kolenati, 1856: 131. Vespertilio serotinus Schreber. 

Amblyotus Kolenati, 1858: 252. Amblyotus atratus Kolenati = Vespertilio nilssonii Keyserling & Blasius. 

PachyomusGray, 1866: 90. Scotophilus pachyomus Tomes. 

Nyctiptenus Fitzinger, 1870: 424. Vespertilio smithii Wagner = Vespertilio hottentota A. Smith. 

Rhinopterus Miller, 1906: 85. Glauconycteris floweri De Winton. Valid as a subgenus. 

Scabrifer Allen, 1908: 46. Substitute for Rhinopterus Allen, thought preoccupied by Rhinoptera Kuhl, 1841, 

Pisces. 

Pareptesicus Bianchi, 1917:lxxvii. VesperugopachyotisDobson. 
Rhyneptesicus Bianchi, 1917: Ixxvii. Vesperugo nasutus Dobson. 
Rhineptesicus Horacek & Hanak, 1985-1986: 16. Lapsus. 

Baculum more or less triangular, its apex occasionally extended into a short shaft, basally rather 
wide, sometimes base expanded into small lobes, tip not expanded, usually more or less pointed or 
gently rounded. There is little flexion in the vertical plane and the tip is not depressed ventrally; 
transversely the base is sometimes slightly arcuate. Externally and cranially not essentially different 
from Pipistrellus but pm 2 invariably absent, the premolar formula being ^ 5 f. 

The karyological features of Eptesicus are summarised by Heller & Volleth (1984) and Zima & 
Horacek (1985). Such as have been examined (andinus, bottae, brasiliensis, diminutus , furinalis , 
fuscus, guadeloupensis , hottentotus, japonensis, lynni, nilssonii, parvus, serotinus, turcomanus) are 
homogeneous in this respect, with 2N = 50, FN = 48-50. On present published knowledge only 
capensis differs with 2N = 32, FN = 50. It is transferred to Pipistrellus by Heller & Volleth (loc. cit.) 
on this account and in the present paper, with others, on bacular grounds. Pipistrellus societatis in 
which 2N is also apparently 50 and FN 48 is transferred to Eptesicus by Heller & Volleth (loc. cit.) 
on account of its karyology and bacular structure, these authors considering it conspecific with P. 
circumdatus (but see Hill & Francis, 1984). Both species are here retained in Pipistrellus, with the 
closely related P. cuprosus. 

Subgenus Eptesicus (Eptesicus) 

Postorbital region not widened, evident postorbital constriction; rostrum not especially shortened, 
its dorsal margins not sharply angular; cranial profile straight or slightly concave, not elevated over 
frontal region; maxillary toothrows almost straight, only slightly convergent; upper surface of 
forearm, tibia and tail lacking horny excrescences. 

(a) nilssonii group 

Cranially large, the skull rather elongate; braincase flattened, elongate, no cranial crests; post- 
orbital region slightly widened; supraorbital area unwidened but with very small supraorbital 
projections; margins of supraorbital region almost straight from postorbital constriction to front 
of orbit, no prominent supraorbital ridges delimiting upper surface of rostrum; the rostrum long, 
not widened, rounded dorsally, its upper surface not flattened but transversely convex above; a 
shallow median rostral depression; slight lateral rostral depressions on each side just above front 
of orbit; cranial profile straight or almost straight, slightly concave over supraorbital region; 
premaxillae not shortened; zygomata moderate with slight jugal process; palate long, narrow, 
interdental palate longer than wide; maxillary toothrows parallel or only slightly convergent; very 
short bony post-palate; prominent basial pits; tympanic bullae not enlarged, not completely cover- 
ing cochleae; i 2 bicuspid, posterior cusp about 1/2-3/4 height of anterior cusp; i 3 wide, as large or 
larger than i 2 in crown area, almost reaching tip of posterior cusp of inner tooth, with very small 
lateral accessory cusps, not displaced anteriorly, lying postero-laterally to i 2 and separated from c 1 
by a modei ate diastema; m 3 not greatly reduced, with trace of fourth commissure, the tooth quite 



252 J. E. HILL & D. L. HARRISON 

long; i t _ 3 slightly imbricated, i 3 a little the largest; pm 2 about 1/2 the crown area and height of 
pm 4 , not compressed in toothrow. 

Included taxa: bobrinskoi (Fig. \3e),gobiensis, nilssonii (Fig. 15a). 

If subgeneric recognition is thought justified for this group then Amblyotus Kolenati, 1858 is 
available. The nilssonii group was recognised as subgenerically valid by Tate (1942a) who however 
included within it a number of taxa here allocated to the nasutus group (vide infra). Strelkov (1986) 
illustrated the bacula of nilssonii, bobrinskoi and gobiensis, considering the last to be a valid species. 

(b) nasutus group 

Cranially small, the skull not especially elongate, braincase flattened, only slightly elongate, 
broad; postorbital region relatively narrower than in nilssonii group; supraorbital area slightly 
widened; margins of supraorbital region nearly straight from postorbital constriction to front of 
orbit, supraorbital ridges sometimes prominent; rostrum shortened, its upper margins slightly 
angular, its upper surface flattened dorso-ventrally, transversely flat, not convex as in nilssonii 
group; a shallow or sometimes more pronounced median rostral depression, slight lateral rostral 
depressions above front of orbit, small lateral rostral elevations above c 1 " ^cranial profile straight 
or nearly so, sometimes slightly concave above supraorbital region; premaxillae sometimes slightly 
shortened; zygomata moderate, on occasion a slight jugal eminence; interdental palate longer than 
wide; maxillary toothrows slightly convergent; short bony post-palate; no basial pits; tympanic 
bullae very large, completely covering cochleae; i 2 large, unicuspid; i 3 small, about 1/2 crown area 
and height of i 2 , with larger main cusp and smaller lateral accessory cusps, anteriorly displaced to 
lie alongside or almost alongside the inner tooth, separated from c 1 by a moderate diastema; m 3 
sometimes reduced, usually with three commissures, no trace of the fourth, antero-posteriorly 
rather short, compressed, platelet-like; i^ _ 3 moderately or well imbricated, similar in size or with i 3 
slightly the largest; pm 2 very small, 1/3-1/4 the crown area and 1/2-1/3 the height of that tooth, 
compressed in toothrow. 

Included taxa: batinensis, matschiei, nasutus (Fig. I4c), pellucens, walli. 

Tate (1942a) included walli, matschiei and pellucens in the nilssonii group but these agree more 
appropriately with nasutus as De Blase (1980) and Honacki et al. (1982) recognised: Ellerman & 
Morrison-Scott (1951) listed matschiei and pellucens as subspecies of nasutus. Indeed, Tate (loc. 
cit.) noted the large tympanic bullae of walli and the absence of basial pits from this taxon. 
Rhyneptesicus Bianchi, 1917 is available if subgeneric recognition is thought justified for this 
group. 

(c) serotinus group 

(c) (i) serotinus subgroup. Cranially large, the skull elongate; braincase flattened, elongate, often 
with lambdoid and sagittal crests forming a distinct occipital 'helmet'; postorbital region slightly 
widened; supraorbital area not widened or only slightly so, with well developed supraorbital ridges 
in many instances; rostrum long, not broadened, its upper surface flattened but less so than in 
nasutus group; very shallow or shallow median frontal depression, shallow to moderate lateral 
frontal depressions just above front of orbit; cranial profile almost straight, a slight concavity 
above front of orbits; premaxillae sometimes a little shortened; zygomata usually robust with 
moderate jugal projection, on occasion slender to moderate, the projection lacking; palate long 
and narrow, the interdental palate longer than wide; maxillary toothrows slightly convergent; 
short bony post-palate; shallow basial pits; tympanic bullae not covering cochleae; i 2 bicuspid, 
posterior cusp about 3/4 height of anterior cusp; i 3 small to very small, 1/2-1/4 or less the crown 
area and height of i 2 , its tip 1/2 or less the height of the posterior cusp of that tooth, with very small 
lateral accessory cusps, the tooth displaced anteriorly to lie alongside or almost alongside i 2 , 
separated from c 1 by a moderate to small diastema, sometimes almost in contact with that tooth; 
m 3 sometimes much reduced, its third commissure obsolescent or obsolete, its second commissure 
short, the tooth platelet-like; ^.3 often massive, much imbricated, i 3 the largest; pm 2 about 
1/3-1/2 the crown area and 1/2 the height of pm 4 . 
Included taxa: andinus (Fig. 13d), argentinus, bottae, brasiliensis (Fig. 13k), dorianus, fidelis, 



VESPERTILIONINE SYSTEMATICS 253 

furinalis (Fig. \3c),fuscus (Fig. 13a), hingstoni, hispaniolae (Fig. 13i), hottentotus, inca, innesi(Fig. 
13j), innoxius, isabellinus (Fig. 13h), megalurus (Fig. 13b), melanopterus , montosus, omanensis 
(Fig. \4a),pachyomus,peninsulae,platyops,punicus, serotinus (Fig. 13g), shirazensis, sodalis, tatei, 
turcomanus. 

Tate (19420) has pointed out that the Old World members of this subgroup fall into two 
categories, one of larger taxa, the other of smaller members of the subgroup. This is also true of the 
New World taxa: however, here the larger forms are found chiefly in North America, extending 
only slightly into South America to which the smaller taxa are entirely confined (Thomas, 1920). 
Material available to us is quite inadequate to attempt any detailed revision and we have followed 
the lead provided by Tate (loc. cit.) in our allocation of all to the one category. Cranial differences 
between large and small members of the subgroup appear chiefly to be those associated with size. 

The subgroup does not extend substantially into Africa. It is represented in Egypt by Eptesicus 
bottae (innesi) and in northwestern Africa by E. serotinus (isabellinus}. Ibanez & Valverde (1985) 
consider the West African platyops to be a subspecies of serotinus, as may be the South African 
hottentotus and also loveni from Kenya. 

(c) (ii) demissus subgroup. Eptesicus demissus Thomas, 1916 from Thailand appears to be known 
only from the holotype, which has a damaged skull. It is very similar to the larger members of the 
serotinus subgroup but has a long bony post-palate, prominent basial pits, i 3 about the same in 
crown area as i 2 and about 1/2 its height, m 3 not especially reduced, its third commissure complete, 
and with i 1 _ 3 moderately imbricated, i 3 the largest. We follow Tate (19420) in referring it to a 
separate subgroup although is likely that more adequate material might enable its status to be 
determined more precisely. 

(c) (iii) (?) pachyotis subgroup. We have been unable to examine Eptesicus pachyotis (Dobson, 
1871) from Assam. Little is known of the species, of which the holotype is in the Indian Museum, 
Calcutta, and as Tate (19420) pointed out, most of the characters given by Dobson in the original 
description might apply to almost any species of Eptesicus. The generic epithet Pareptesicus 
Bianchi, 1917 was proposed for this taxon. 

Subgenus Eptesicus (Rhinopterus) 

Cranially small; braincase low, flattened and elongate, inflated anteriorly; postorbital region wide; 
supraorbital area widened with very small supraorbital tubercles; anterior margin of orbit flange- 
like; rostrum short, flattened dorso-ventrally, its dorsal margins angular; very shallow median 
rostral depression, shallow lateral depressions just above front of orbit; cranial profile convex, 
raised above frontal region; premaxillae not shortened; zygomata slender, no jugal projection; 
palate short, broad, interdental palate about as long as wide; maxillary toothrows convergent; 
short to moderate bony post-palate; no basial pits; i 2 bicuspid, posterior cusp 3/4 or more the 
height of the anterior cusp; i 3 small or minute, about 1/4 or less the crown area of i 2 , about 1/3-1/2 
its height, its lateral accessory cusps very small or obsolete, lying postero-laterally or almost 
alongside the inner tooth, separated from c 1 by a moderate or small diastema; m 3 not much 
reduced, its third commissure complete; i l _ 3 strongly imbricated, i 2 the smallest, i l and i 3 of 
similar size; pm 2 very small, about 1/2 crown area and height of pm 4 , strongly compressed in row; 
horny excrescences on upper surface of forearm, tibia and tail. 
Included taxa.:floweri (Fig. 13f), lowei(Fig. 131). 

The status of the 'NycticeinF 

An especially interesting feature emerges from our survey of bacular morphology in the Vesperti- 
lioninae. The structure of the baculum suggests very strongly that the 'Nycticeini' (or 'Nycticeiini') 
as presently accepted is not a natural group. Defined chiefly on dental characters (i 2 generally 
unicuspid, i 3 and pm 2 absent), this group was assembled by Tate (19420) to include Baeodon, 
Rhogeessa, Otonycteris, Nycticeius (i.e. N. humeralis, including cubanus), 'Scoteinus' (then includ- 
ing among others the Australian species now referred to Scoteanax and Scotorepens), Scotoecus, 
Scotomanes and Scotophilus. Tate, however, made no mention of the African species schlieffenii 



254 J. E. HILL & D. L. HARRISON 

which was extralimital to his study but which by then had been variously referred either to 
Scoteinus (Miller, 1907) or to Nycticeius (Hollister, 1918; Braestrup, 1935). More recently, the 
type species of Scoteinus (the Indian emarginatus) has proved to be a Scotomanes (Sinha & 
Chakraborty, 1971) and the other Indian species (pallidus) formerly referred to it a Scotoecus 
(Hill, 1974). The Australian Scoteanax and Scotorepens have been considered to be subgenera of 
Nycticeius (Laurie & Hill, 1954; Koopman, 1978; Corbet & Hill, 1980) but recently have been 
accorded generic rank (Kitchener & Caputi, 1984; Corbet & Hill, 1986). Thus the current concept 
of Nycticeius is of two species, N. humeralis from North America and TV. schlieffenii from Africa 
and southwestern Arabia. 

The bacula ofRhogeessa (Fig. 1 8k) and Baeodon (Fig. 1 5b) are quite distinctive and are variants 
of the saddle-like or slipper-like structure found in Myotis and Plecotus or their allies, as are the 
very characteristic bacula of Otonycteris (Fig. 16a) and Nycticeius humeralis (Fig. 17k), the type 
species of Nycticeius. Scotomanes (Fig. 18g) and Scotophilus (Fig. 17g-j) have bacula reminiscent 
of the flattened, triangular structure of Eptesicus and its immediate associates. In contrast, the 
bacula of Scotoecus (Fig. 20a-e), Nycticeius schlieffenii (Fig. 16e), Scoteanax (Fig. 16i), and 
Scotorepens (Figs 16g, h, 21e, f) are closely similar to those of Pipistrellus (Pipistrellus). Thus in 
bacular terms this supposed group appears to be a composite of different elements, so dissimilar 
among themselves that its unity seems very unlikely. Kitchener & Caputi (1984) contended on 
the grounds of a phyletic analysis that Otonycteris and Scotophilus nigrita fitted poorly into the 
then current concept of the 'Nycticeini' and moreover on similar considerations that Nycticeius 
humeralis and Nycticeius schlieffenii are not congeneric. This view contrasts sharply with that of 
Koopman ( 1 978) who remarked that the latter are similar in all important characters and should be 
retained together in the subgenus Nycticeius (Nycticeius). 

The sharp bacular difference between the American humeralis and the African schlieffenii 
suggests wider separation and indicates that their congeneric association is wrong, despite their 
morphological similarities in some other ways. We propose therefore to dissociate schlieffenii from 
Nycticeius as generically distinct. The newly proposed genus may be called: 



Nycticeinops gen. nov. 

TYPE SPECIES: Nycticejus schlieffenii Peters, 1 860. 
REFERRED SPECIES: None. 
DISTRIBUTION: Mauretania to Egypt, Namibia and Mozambique; SW Arabia. 

DIAGNOSIS: Baculum (Fig. 16e) distinctive, with expanded base and long fluted shaft, very different 
from that of Nycticeius humeralis (Fig. 17k) which is slipper-like, elevated proximally and distally; 
cranially similar to Nycticeius sensu stricto but rostrum shorter, more narrowed anteriorly, the 
maxillary toothrows much more convergent, not nearly parallel, with correspondingly narrower 
narial and anterior palatal emarginations; mandible similarly narrowed anteriorly, with i 1 _ 3 
strongly imbricated, thrust further anteriorly into an arc; narial emargination more clearly 
U-shaped, not prolonged posteriorly; anterior palatal emargination extending further posteriorly; 
basial depressions absent or only very slight; pm 2 more reduced. Similar to Australian Scoteanax 
and Scotorepens but differing sharply in bacular morphology, the baculum with a more flanged and 
fluted shaft and lacking the modification of the tip found in these genera, and in less reduction of 
mf 

Differs from Pipistrellus in massive, unicuspid i 2 which has no trace of a secondary cusp, in 
contact or nearly so with c 1 , the premaxillae greatly shortened, combined with the almost invari- 
able absence of i 3 and pm 2 . Similar in some respects to Scotozous but differing in bacular 
morphology; in the presence of a small, posteriorly directed lobe at the base of the inner margin of 
the ear; tip of tragus anteriorly directed; pm 2 almost invariably absent; pm 2 more reduced. Similar 
also in some ways to Scotoecus but penis not greatly lengthened, baculum similarly shorter, its tip 
not expanded and bifid; rostrum narrower, uninflated; narial and anterior palatal emarginations 
not extensively deepened; and anterior face of c 1 rounded, not flattened and grooved. 



VESPERTILIONINE SYSTEM ATICS 255 

DESCRIPTION: Small (length of forearm about 29-33 mm); muzzle flattened, anteriorly sparsely 
haired, nares opening obliquely; ear rounded with broadly rounded tip, anterior or medial margin 
with small, posteriorly directed basal lobe, anterior margin slightly convex for most of its length; 
posterior margin nearly straight distally, more convex proximally with well developed, thickened 
antitragal lobe; tragus with bluntly pointed, anteriorly directed tip, the anterior margin strongly 
concave basally, straight distally, upper margin nearly horizontal, posterior margin strongly con- 
vex, with prominent basal lobe; calcar extending along a little more than one half of the uropatagial 
border; well developed, rounded post-calacarial lobe or epiblema. Dorsal surface of head and body 
brown to pale brown, the pelage unicolored; ventral surface paler brown to greyish white, the 
pelage usually unicolored but in the darker subspecies faintly bicolored, the hair bases darker than 
the tips. 

Skull low, with broad, flattened braincase, not elevated frontally; very low cranial crests and 
very slight occipital 'helmet'; postorbital region wide; supraorbital area a little broadened; rostrum 
not expanded laterally, narrow anteriorly; cranial profile almost straight, a little depressed over 
front of orbits; narial emargination U-shaped, extending posteriorly one half of distance from tip 
of maxillae to a line joining front of orbits; premaxillae much shortened; anteorbital foramen 
moderate to large; zygomata slender, no jugal projection; palate rather short, the interdental palate 
little longer than wide, narrowed anteriorly, maxillary toothrows convergent, anterior palatal 
emargination narrow, extending posteriorly to a line joining the posterior faces of c 1 " 1 , not 
extending laterally beyond the inner faces of i 2 " 2 ; short to moderate bony post-palate, a narrow 
median post-palatal spine; basial depressions at best only very slight. 

Dental formula normally if 23, prnE^l m| 2 f| = 30. Upper incisor i 2 massive, unicuspid, usually 
separated from c 1 by a very short diastema, sometimes in contact with this tooth; i 3 and pm 2 almost 
invariably absent (Thomas, 1 890; Thomas & Wroughton, 1 908); pm 4 in contact with c 1 , with small 
protocone; lingual shelves of m 1 ~ 3 widely separated, m 3 not reduced, with three commissures and 
mesostyle, about 1/2 crown area of m 1 or m 2 ; ^ _ 3 strongly imbricated to one half of their width, 
thrust forward, i l clearly tricuspid, i 2 _ 3 less obviously so, i ^ longest, i 2 _ 3 more massive; pm 2 much 
reduced, 1/2-1/4 crown area of pm 4 and 1/2 its height, compressed in toothrow; m 3 slightly 
reduced, posterior triangle smaller than anterior triangle, hypoconid and entoconid lower than 
protoconid, paraconid and metaconid. 

Thomas & Wroughton (1908) reported a specimen (BM(NH) 8.4.3.23) from Tette, Malawi in 
which a well developed i 3 is present in the left side of the jaw. Dobson (1878) remarked of two 
specimens in the Museum National d'Histoire Naturelle, Paris that pm 2 is present on one side in 
one, on both sides in the other, but Thomas (1890) who examined these noted that pm 2 is com- 
pletely absent from one and present on both sides in the other, Dobson having in the first instance 
perhaps mistaken a grain of sand for the tooth. Allen (1914) remarked of a specimen that he 
identified as schlieffenii from Bados, Blue Nile Province, Sudan that pm 2 was present on both sides 
of the jaw and that 'in common with Scotoecus, it has a large penial bone, 12mm long' but 
Koopman (1965) pointed out that in fact this specimen is a Scotoecus (not Scotophilus as 
Qumsiyeh, 1985 avers) and that schlieffenii has a very much smaller penis. 

INCLUDED TAXA: The genus is monospecific, its sole species N. schlieffenii Peters, 1860. Taxa 
allocated to it either as valid subspecies or synonyms include adovanus Heuglin, 1877; africanus 
Allen, 1911; albiventer Thomas & Wroughton, 1908; australis Thomas & Wroughton, 1908; 
bedouin Thomas & Wroughton, 1908; cinnamomeus Wettstein, \9\6\fitzsimmonsi Roberts, 1932; 
minimus Noack, 1887. 

ETYMOLOGY: The name of the new genus is derived from vu, VUKTOCT or vuKiioa, night, and oy 
aspect. 

REMARKS: The type species schlieffenii has undergone a wide variety of generic allocations and 
taxonomic change since Peters ( 1 860) first described it as a Nycticejus. Dobson ( 1 876, 1 878) placed 
it in Scotozous with dormeri while under the impression that this genus lacked i 3 , and considered 
(1878) Scotozous to be a subgenus of Vesperugo. Noack (1877) and Thomas (1890) referred it to 
Scotophilus, the latter author discussing this genus in relation to Scotozous, which following 



256 J. E. HILL & D. L. HARRISON 

Dobson he thought to have but one pair of upper incisors. Trouessart (1897) initially followed 
Dobs~>n (1878) in allocating schlieffenii to Scotozous as a subgenus of Vesperugo, but later (1904) 
changed this opinion to consider Scotozous a subgenus of Scotophilus. Miller (1907) referred 
schlieffenii to Scoteinus, although in fact the species does not display the reduction of m| that he 
considered diagnostic for this genus and which occurs in the Australian species (balstoni, greyii, 
now incorporated into Scotorepens) that he allocated to it. Miller's view was adopted by Thomas & 
Wroughton (1908) and in differing ways by many subsequent authors. However, Allen (1911) 
when describing africanus referred it to the hitherto American genus Nycticeius, commenting on its 
similarity to N. humeralis and Hollister (1918) remarked that Old World bats usually placed in the 
genus Scoteinus did not seem to differ generically from the American species of Nycticeius, to which 
he also referred africanus. Since then africanus has been relegated to subspecific status or synonymy 
in schlieffenii (Braestrup, 1935; Allen, 1939; Aellen, 1952). Braestrup (loc. cit.) also employed 
Nycticeius for schlieffenii in preference to Scoteinus, and pointed out that its last upper molar was 
not reduced in the way that Miller (1907) had described for that genus. This author drew attention 
to the affinity thus established between the Ethiopian and American faunas, but did not exclude the 
possibility of convergent evolution from different Pipistrellus-\ike forms. Tate (1942a) maintained 
Nycticeius and Scoteinus as distinct genera but Simpson (1945) united them, a lead followed by 
many modern authors who have considered Scoteinus a subgenus of Nycticeius. Thus Ellerman & 
Morrison-Scott (1951) and Ellerman et al. (1953) referred schlieffeni to Scoteinus as a subgenus 
of Nycticeius, while Laurie & Hill (1954) listed the Australian species before then allocated to 
Scoteinus in Scoteanax and Scotorepens as further valid subgenera of Nycticeius. On the other 
hand, Rosevear (1965) considered Nycticeius and Scoteinus synonymous. Koopman (1965) 
referred schlieffenii to Scoteinus as a subgenus of Nycticeius but later (in litt. in Hayman & Hill, 
1971) revised this opinion to allocate it to Nycticeius (Nycticeius), since then (1978) reinforcing this 
view. 



The classification of the Vespertilioninae 

Earlier classifications of the Vespertilioninae (Miller, 1907; Tate, 19420) rely heavily on the pattern 
of reduction of the incisor and premolar teeth, chiefly on the presence or absence of the outer upper 
incisor (i 3 ), of one or both of the first (pm 2 ) or second (pm 3 ) upper premolars, and on the presence 
or absence of the second (pm 3 ) of the lower premolars, as Tate's 'phyletic' diagrams (loc. cit.) 
indicate. These dental features have been discussed in more detail above (p. 230): they reflect 
the degree of shortening that forms an evident trend within the subfamily. When combined with 
the relative size of one or more of these teeth and the position of the relevant tooth or teeth in the 
toothrow such factors form an important element in generic identification and diagnosis (cf. 
Miller, loc. cit.). The many different combinations of incisive and premolar formula in the sub- 
family (Table 2), the evanescence in some genera of some of the teeth involved, the extreme 
tendencies towards reduction seen in some such as Pipistrellus, and the variety of positions within 
the toothrow adopted by i 3 and pm 2 in particular reinforce the conclusion that such features reflect 
a universal trend that may have occurred more than once within the group and which as a result 
may not provide a totally reliable yardstick by which relationship may be judged. 

In addition to these dental features, Tate (1942a) reviewed a number of other characters used in 
the classification of the subfamily. These include the presence or absence of accessory canine cusps; 
the form and shape of the braincase and rostrum; the degree of reduction of the zygomata; the 
structure of the palate, its anterior emargination and accessory anterior and posterior spines; the 
presence or absence of basial pits; enlargement of the ears and their associated bony structures; 
the presence or absence of adhesive pads on the thumb or foot; and the nature of other minor 
structures such as the calcar. These features, however, seem of greater value in the distinction of 
species and species groups, that is, for infrageneric classification, or for the diagnosis of individual 
genera. 

The value of such characters has been discussed at some length by Zima & Horacek (1985) 
who pointed out that there are grounds for thinking that some of the traditional morphological 



VESPERTILIONINE SYSTEMATICS 257 

characters may not provide unequivocally reliable criteria for the establishment of a classification 
based on presumed phyletic relationship, and that their taxonomic significance may be limited. 
They also remarked that such characters may reflect parallelism or convergence, or result from 
selection pressure rather than relationship. These reasons led them to suggest that the baculum 
might provide one of several alternative sources of reliable, taxonomically useful criteria based on 
characters that do not have a direct adaptive significance. 

The structure of the baculum in the Vespertilioninae suggests some modifications to tribal 
classification within the subfamily, although clearly other morphological characters need to be 
given equivalent or greater weight. Provisionally, therefore, we offer an arrangement of the Vesper- 
tilioninae in which bacular morphology is used in association with the traditional diagnostic 
features to suggest possible relationship. This classification is presented in Table 1 . 

There appear to be two major bacular types in the Vespertilioninae, each with numerous varia- 
tions as might be expected in such a large and diverse subfamily. A classification that includes a 
major consideration of bacular morphology shows significant resemblances to earlier arrange- 
ments based on traditional and conventional morphological features. However, there are some 
wide divergences, as for example the seemingly artificial nature of the 'Nycticeini' or the associ- 
ations of the various genera of big-eared bats. Tate (19420) commented upon the latter and 
pointed out that very large ears and their associated auditory specialisations in the skull occurred 
independently in three sections of the subfamily: indeed, if Antrozous and Bauerus are included, 
these features occur four times in the group. In particular, both Miller (1907) and Tate (loc. cit.) 
associated Laephotis with Histiotus on cranial and dental morphology but its bacular structure 
shows a clear affinity with Pipistrellus (Neoromicid) as here recognised. Otonycteris, another 
big-eared bat, was allied by Tate (loc. cit.) to the 'Nycticeini' but proves to have a baculum much 
more like those of the plecotine genera. 

One major bacular type is 'saddle-like' or 'slipper-like' and is exemplified by Myotis and 
Pizonyx. Their bacula are very similar, emphasising the close relationship that is generally accepted 
between these genera. The baculum of Lasionycteris is somewhat different in the presence of a 
lengthened shaft. However, in comparison with the long-shafted bacula found in the Pipistrellini 
the baculum of Lasionycteris is relatively short, and it retains indications of the more characteristic 
myotine type in its upraised proximal and distal portions. The occasional presence of a flattened 
dorsal prominence on its base also recalls the condition found in Idionycteris. The genus, although 
having some specialised features, is allied firmly to Myotis by Miller (1907) and Tate (\942a). It has 
slightly hooked upper incisors, i 3 with a slightly caniniform profile as in Myotis', pm 2 is in the line of 
the toothrow; m 3 is unreduced; pm 2 _ 3 are exactly as in Myotis, much smaller than pm 4 , with pm 3 
not removed from the line of the other teeth. Although pm 3 has been lost, this appears to be a 
specialisation; as Tate (loc. cit.) pointed out, pm 2 _ 3 still agree closely with those of the less 
specialised species of Myotis not only in relation to each other but also in their proportional size 
relative to pm 4 . Although associated with Myotis, this genus is considerably specialised in other 
ways (Miller, loc. cit.) and its bacular structure may well reflect this divergence. Its baculum might 
be regarded as derived from the more typical myotine structure. 

Bacula variously reminiscent of the saddle shaped structure found in Myotis occur in a number 
of other genera. Such bacula characterise Plecotus (including Coryhorhinus), Idionycteris, 
Barbastella, Rhogeessa, Baeodon, Nycticeius, Otonycteris, Lasiurus, Dasypterus, Antrozous and 
Bauerus, and possibly may be found in Euderma. Tate (19420) postulated the grouping 'Plecotini' 
for Plecotus, (Corynorhinus), Idionycteris and Euderma, allying it to the Myotini but not employing 
the term in a formal taxonomic or systematic sense. Bacular morphology thus lends support to his 
hypothesis that the plecotine genera should be associated with Myotis. Also, the baculum of 
Barbastella suggests that it too belongs here: Miller (1907) postulated such a relationship, despite 
several morphological differences. Rhogeessa, Baeodon, Nycticeius and Otonycteris also seem 
allied to this grouping. Tate (19420) referred these genera to the 'Nycticeini' with Scotoecus, 
Scotomanes and Scotophilus on account of their incisive and premolar dentition. However, the 
bacula of Rhogeessa, Baeodon, Nycticeius and Otonycteris are variants of the saddle-like type; that 
of Scotoecus is like that of Pipistrellus (Pipistrellus), and the bacula of Scotomanes and Scotophilus 
are broadly similar to those ofEptesicus and its allies. Lasiurus, Dasypterus, Antrozous and Bauerus 



258 J. E. HILL & D. L. HARRISON 

have further variants of this bacular type, but are quite distinctive on other morphological 
grounds. 

The bacula of Antrozous and Bauerus are not at all like that of Otonycteris, with which these 
genera have been tentatively associated (Pine et al., 1971), nor do their bacula have any significant 
resemblance to those of Nyctophilus or Pharotis, thus supporting the view (Koopman, 19846, 
1985; Breed & Inns, 1985) that these North American genera should not be associated with 
the Australian Nyctophilus and Pharotis in the subfamily Nyctophilinae. Bacular morphology 
suggests instead an association with those genera that have the myotine type of baculum, to which 
the bacula of Antrozous and Bauerus have many resemblances. The bacula of Nyctophilus and 
Pharotis (Fig. 22a-h) are consistently homogeneous and differ in many ways from those of the 
genera usually referred to the Vespertilioninae. For the present we would place these two genera in 
a separate subfamily, the Nyctophilinae, rather than merge them into the Vespertilioninae as is 
done by Koopman (1984a, 19846, 1985). 

A further basically triangular and flattened variant of the saddle-like baculum characterises the 
genera Eptesicus, Vesper tilio (if the pseudobaculum is ignored), la and Histiotus. Miller (1907) 
remarked that the skull of Vespertilio showed a strong likeness to that of Lasionycteris but that 
the former was in all respects a typical Eptesicus. Vespertilio and Lasionycteris are separated 
by marked dental and bacular differences: the bacular morphology of Vespertilio allies it with 
Eptesicus as Miller suggested. It is perhaps not unreasonable to speculate that Lasionycteris which 
has a strongly myotine dentition has diverged among the Myotini in the same way as Vespertilio 
has diverged among the Vespertilionini, the latter genus supporting a long penis either by a 
centrally situated baculum or perhaps more effectively by the development of a cartilaginous 
pseudobaculum, this function in Lasionycteris by a short shaft. The genera Tylonycteris and 
Mimetillus also belong here. The African Glauconycteris has been associated (Ryan, 1966; 
Koopman, 1971) with the Australian Chalinolobus but their bacula differ widely. Although struc- 
turally variable within the genus, the bacula of Glauconycteris are more like the vespertilionine 
or eptesicine type: those of Chalinolobus are long-shafted and like the bacula of Pipistrellus 
(Pipistrellus). Finally, the baculum ofScotomanes appears to be a derivative of the saddle-like type, 
leading to the distinctive baculum of Scotophilus. 

The genus Pipistrellus seems to stand more or less at the centre of the second major grouping. 
It has broadly two divisions in bacular terms, one characterised by a long baculum with well 
developed basal lobes and a relatively long, mostly cylindrical shaft, its tip often bifid or with 
similar elaboration. The second division includes those species in which the basal lobes are some- 
times small or obsolete and which have a shorter, flatter, ventrally fluted shaft, its tip sometimes 
elaborated into a spatulate or platelet-like structure. 

These groupings have been used in this study to support subgeneric division of this large genus. 
The first division includes Pipistrellus (Pipistrellus}, P. (Vespadelus), P. (Perimyotis} and P. 
(Arielulus}. Reduction and loss of pm 2 occurs in P. (Arielulus) and the tooth is almost invari- 
ably absent in P. (Vespadelus}. The second division contains P. (Hypsugo) in which pm 2 may be 
very small or absent, P. (Neoromicia) from which it is again almost invariably absent, and P. 
(Falsistrellus}. Although primarily Old World in distribution, both of these divisions are repre- 
sented in the New World, each by a single species. The Australian P. ( Vespadelus} seems on bacular 
features to represent P. (Pipistrellus); the wholly African P. (Neoromicia) is apparently similarly 
related to P. (Hypsugo), of which P. (Falsistrellus} appears to be an eastern representative. 

The majority of the genera here allocated to the Pipistrellini show strong bacular affinities to 
Pipistrellus (Pipistrellus): some such as Glischropus and Scotozous have been considered congeneric 
with Pipistrellus in the past. Besides Glischropus and Scotozous these include Nycticeinops, 
Scoteanax, Scotorepens, Scotoecus, Nyctalus, Hesperoptenus and Chalinolobus, all with long- 
shafted bacula. Of the remainder, Laephotis in bacular structure is similar to P. (Neoromicia}, while 
Philetor has a baculum that appears to be an elaboration of the bacular structure found in some of 
P. (Hypsugo). Tate (1942a) postulated a relationship between Philetor, Tylonycteris and perhaps 
Mimetillus but the bacula of the first two are totally dissimilar and the structure is apparently 
absent from Mimetillus: it is very small in Tylonycteris. Hill (1966a) drew attention to the unusual 
genitalia of Philetor and following Tate's (loc. cit.) suggestion of affinity with Pipistrellus joffrei and 



VESPERTILIONINE SYSTEMATICS 259 

its associates allied Philetor with this group. Unfortunately, excepting for the aberrant species 
stenopterus the bacula of the stenopterus subgroup (including P.joffrei) of this present study remain 
unknown. 

Bacular morphology suggests that the conventional view that Eptesicus and its immediate allies 
derive from or are closely related to Pipistrellus can be questioned. Cranially and dentally there are 
many similarities between 'Eptesicus ' as formerly denned and Pipistrellus and as Koopman (1975) 
has pointed out, the loss of pm 2 enables a species to cross the boundary between the two genera as 
then understood, a process which in his view might have occurred more than once. Our conclusions 
do not challenge this opinion: those 'Eptesicus' species in which pm 2 has been found occasionally 
to occur prove on bacular grounds to be closer to Pipistrellus than to Eptesicus as we understand it, 
while Pipistrellus as formerly defined has long been known to include some species from which on 
occasion this 'diagnostic' tooth is absent. Clearly, our findings support Koopman's (loc. cit.) 
opinion that this process may have occurred several times and indeed may be occurring in some 
species, but all belong to the one genus, Pipistrellus. 

As we understand its composition, Eptesicus is now a more restricted genus in which the tri- 
angular, flattened baculum is basically closer in structure to the saddle-like grouping than to the 
long-shafted group, although some Eptesicus do indeed have bacula that suggest the beginnings 
of basal lobulation or of a very short shaft. We suggest therefore that in bacular terms the 
Vespertilionini to which we refer Eptesicus may represent a transitional stage between the saddle- 
like baculum and the predominantly basally lobed and long-shafted type. Tylonycteris and 
Glauconycteris also show this tendency. 

Dental reduction proceeds throughout both of the major bacular groups. In the grouping with 
broadly myotine or saddle-like bacula the dentition varies in number of teeth from a total of 38 
(Myotis, Pizonyx) through 36 (Lasionycteris, Plecotus and allies), 34 (Barbastella, Eptesicus and 
allies), 32 (Lasiurus), 30 (Dasypterus, Rhogeessa, Baeodon, Nycticeius, Otonycteris, Scotomanes, 
Scotophilus) to 28 (Antrozous, Bauerus}. In the second of the two major bacular groups, dental 
reduction varies from Eudiscopus with a total of 36 teeth (its association here is presumed) 
through 34 (Pipistrellus, Glischropus, Scotozous, Nyctalus, Chalinolobus), 32 (Laephotis, Philetor, 
Hesperoptenus) to 30 (Nycticeinops, Scoteanax, Scotorepens, Scotoecus). Thus this trend occurs 
concurrently in the two major groupings, taking the same form in each by increasing the size and 
bulk of i 2 , the reduction, transposition and loss of i 3 , and the progressive reduction, transposition 
and loss of pm 2 , pm 3 and pm 2 . 



Zoogeographical considerations 

The saddle-shaped or slipper-like baculum characteristic of the Myotini, Plecotini, Lasiurini and 
Antrozoini as here understood is cosmopolitan in but one genus, Myotis. It occurs in one Holarctic 
genus, Plecotus, in one Palaearctic genus, Barbastella, itself probably closely related to Plecotus, 
and in one other Old World genus, Otonycteris, that occurs in southwestern Asia and northern 
Africa. Otherwise this bacular type is limited to the New World. Lasionycteris, exclusively North 
American, has a baculum apparently derived from this type, as does Nycticeius, also North 
American, although in this genus the baculum is considerably modified to the extent that Hamilton 
(1949) commented upon its unique character among the genera that he had examined. Thus 
although the saddle-shaped baculum or its derivatives is represented about equally in number of 
species in the Old and New Worlds, genera with bacula of this type predominate in the latter, its 
extension into the Old World being primarily through the many species of Myotis, with a lesser 
contribution from Plecotus, Barbastella, and Otonycteris. 

A further variety of this bacular type is found in the Vespertilionini, that is, in Eptesicus and its 
close relatives. In these, the baculum is less strongly saddle-shaped or slipper-like, flatter, and often 
more triangular in outline. This bacular type is primarily Old World in numbers of genera and 
species, only Eptesicus among Old World genera extending to the New World where there is a 
closely related genus, Histiotus. In the Old World, Vespertilio is also closely related to Eptesicus. 
Another Old World genus, la, is a giant representative of this same bacular type. The southeastern 



260 J. E. HILL & D. L. HARRISON 

Asian Tylonycteris and the African Glauconycteris have bacula that are modified variants of this 
type: Mimetillus, in which no baculum has been found, also appears to belong here. Two further 
Old World genera, Scotomanes and Scotophilus, also have bacula that are similar in many respects 
to the vespertilionine type. 

The shafted or long-shafted bacular type is confined almost exclusively to the Old World, and is 
represented in the New World by no more than two species of Pipistrellus in the Nearctic region, 
one of these with a highly modified baculum. This bacular type is restricted to the Pipistrellini and 
within that grouping to those genera that for the most part can be shown on other grounds to 
cluster around Pipistrellus. Indeed, some such as Scotozous, Glischropus, Scoteanax, Scotorepens 
and perhaps even Nyctalus might on bacular grounds be regarded as subgenera of this widespread 
genus. In a reduced form this bacular type appears in two of the subgenera of Pipistrellus, P. 
(Perimyotis) and P. (Arielulus). Widespread in the Palaearctic region and in southeastern Asia, 
this bacular type is represented in Australia by five distinct groupings: Pipistrellus (Pipistrellus}, 
P. ( Vespadelus), Scoteanax, Scotorepens, and Chalinolobus. This type of baculum also occurs in 
Africa among Pipistrellus kuhlii and its associates, which might in fact be considered to warrant 
recognition as a further subgenus of Pipistrellus. 

A further variant of the shafted bacular type is found in Pipistrellus (Hypsugo) and P. (Falsistrel- 
lus). In these the shaft is shorter and is ventrally fluted, often with expansion of the tip. Pipistrellus 
(Hypsugo) is confined chiefly to Asia and Africa, where in the latter region it appears to be closely 
associated with P. (Neoromicia) in which pm 2 is generally lost. Thus as in Australia where P. 
(Vespadelus) in which pm 2 is also generally absent appears to derive from P. (Pipistrellus), so in 
Africa P. (Neoromicia) is apparently similarly related to P. (Hypsugo). Of the two North American 
pipistrelles, P. subflavus has a reduced form of the P. (Pipistrellus) baculum, the shaft very short 
and stubby: this species has a myotine tragus and has been considered (Menu, 1984) to have a 
myotine dentition. However, on the balance of features it appears to be clearly referable to 
Pipistrellus and indeed to be cranially and dentally close to P. (Pipistrellus), which apparently it 
represents in North America. There do not appear to be sufficient grounds to justify its generic 
separation from Pipistrellus as has been recently effected (Menu, loc. cit.), although subgeneric 
recognition within that genus seems appropriate. The second North American species of Pipistrel- 
lus, P. Hesperus, should evidently be referred to P. (Hypsugo) with which it has close bacular and 
dental similarities, although recently generic separation (Horacek & Hanak, 19850, b, 1985-1986) 
has been proposed for it. Finally, P. (Falsistrellus) is restricted to southeastern Asia, Australasia 
and Tasmania: the deeply ventrally fluted baculum of this subgenus, lacking basal and distal 
modification but massive and substantial appears to be an extreme of the P. (Hypsugo) type: 
possibly P. (Falsistrellus) represents P. (Hypsugo) which seems to be linked to it by several of its 
Asian species. 

One corollary of the removal of the African capensis and tenuipinnis groups of 'Eptesicus' to 
Pipistrellus, and of the similar transfer of the Australian species formerly referred to 'Eptesicus' is 
that in the Old World Eptesicus now becomes primarily Palaearctic, with outliers, perhaps all 
closely connected to E. serotinus, in Africa while in the New World it extends over both North and 
South America. In southeastern Asia the genus becomes restricted to no further east than southern 
Thailand, the former enormous hiatus in its distribution between this part of southern Asia and 
Australia having been removed. 



Conclusions 

(1) The current classification of the Vespertilioninae is based chiefly on adaptive characters with 
considerable emphasis on facial shortening and concomitant dental reduction and loss. Several 
authors have drawn attention to the deficiencies and dangers of any classification that relies heavily 
on such features. A review of bacular morphology within the subfamily suggests that this structure 
provides indications of relationship that in many respects support the existing classification but 
which also indicate several changes to the current arrangement. In particular, bacular morphology 
suggests a number of major and minor changes in the systematics of the nominal genera Pipistrellus 



VESPERTILIONINE SYSTEMATICS 261 

and Eptesicus, separated hitherto only by dental formula, itself subject to variation in both 'genera' 
as they are currently understood. 

(2) The presence or absence of the anterior upper premolar (pm 2 ) in Pipistrellus and Eptesicus, 
used formerly as their principal diagnostic character, has little taxonomic significance. The tooth is 
variable in Pipistrellus as here understood, being reduced or lost in three of its subgenera, and is 
absent from Eptesicus as we envisage it. 

(3) Bacular morphology in Pipistrellus and Eptesicus provides groupings that largely agree in 
species content with those proposed by earlier authors such as Tate (19420) and Koopman (1973, 
1975) although in basing their studies on 'conventional' morphological characters neither con- 
sidered these genera in their entirety. The bacular morphology of 'Eptesicus' as it is currently 
understood provides a clear indication that as such it is not a natural group, but that three species 
aggregations, the Australian pumilus group and the African capensis and tenuipinnis groups, 
should be transferred to Pipistrellus. 

(4) It has been possible to recognise and define subgenera for the major species groups in both 
Pipistrellus and Eptesicus and to suggest possible relationships between them. One subgenus is 
described as new as Pipistrellus (Arielulus) for P. circumdatus and its allies. 

(5) There appear to be clear links between certain of the pipistrelline subgenera: Pipistrellus 
( Vespadelus} in Australia seems to represent P. (Pipistrellus) in bacular terms while P. (Hypsugo) is 
apparently represented in Indo-Australia by P. (Falsistrellus) and is related to the African P. 
(Neoromicid). Although the features of the two Nearctic species of Pipistrellus have been thought 
to justify their recognition in separate, individual genera we consider that the characters of one 
(subflavus) merit no more than subgeneric status as the sole species of P. (Perimyotis), which itself 
perhaps represents P. (Pipistrellus), while the other (Hesperus) is perhaps more appropriately 
referred to P. (Hypsugo). 

(6) The examination of bacula in Pipistrellus has suggested that some taxa hitherto ranked as 
subspecies, for example abramus, paterculus or helios, might in fact be distinct species. 

(7) As we now understand the species content of Pipistrellus and Eptesicus the former remains 
primarily an Old World genus where it is widespread and diverse in the tropics and subtropics, 
extending into the temperate zones and just to North America. In contrast, our concept of 
Eptesicus limits this genus to the New World and in the Old World primarily to the Palaearctic, 
with outlying representatives in Africa. It does not extend significantly into Australasia. 

(8) Bacular morphology suggests the informal recognition of two major groupings within the 
subfamily Vespertilioninae. The first includes the Myotini, Plecotini and Lasiurini; Antrozous and 
Baeurus, which in bacular terms have no relation to Nyctophilus and Pharotis (the Nyctophilinae); 
the Scotophilini to include Scotomanes and Scotophilus; and finally the Vespertilionini, here 
reduced in content to include Eptesicus and its close relatives Histiotus, la and Vespertilio, with 
Tylonycteris, Mimetillus and Glauconycteris. 

(9) The second grouping consists of Pipistrellus and those genera which cluster round it. All with 
the possible exception of Philetor appear to relate quite closely in bacular terms to one or other of 
the subgenera that we recognise in Pipistrellus, principally to P. (Pipistrellus). Laephotis, formerly 
considered related to Histiotus, is instead in bacular terms closely associated with P. (Neoromicid). 
The bacula of Chalinolobus and Glauconycteris are widely dissimilar athough these genera have 
been closely allied in the past; Chalinolobus is of the pipistrelline type while the baculum of 
Glauconycteris apparently associates it more appropriately with Eptesicus and its allies. 

(10) Bacular morphology provides clear indications that the 'Nycticeini' of Tate (\942a) and 
Koopman (1984, 1985) is not a natural group, its constituent members despite cranial and dental 
similarities having widely different bacula. Thus Rhogeessa, Baeodon, Nycticeius sensu stricto, and 
Otonycteris have been here allied to the plecotine bats on bacular grounds, while Scoteanax, 
Scotorepens and Scotoecus are quite clearly associates in bacular terms of Pipistrellus. 'Nycticeius', 
at one time thought to include the Australian Scoteanax and Scotorepens as well as its North 



262 J. E. HILL & D. L. HARRISON 

American type species humeralis and the African schlieffenii, has recently been restricted only to the 
American and African forms. These prove to have widely different bacula; humeralis has been 
associated with the plecotine bats on this account, while generic status has been accorded to 
schlieffenii with the proposal of a new generic name, Nycticeinops. 

(11) The two broad bacular types that we discern in the subfamily Vespertilioninae have definite 
geographical patterns: the saddle-like baculum and its variants that characterise the first group 
noted above is primarily New World and Palaearctic, extending less significantly into the Old 
World tropics or Australasia, while the shafted baculum of the second group is chiefly confined to 
the Old World. 



Addendum 

A phenetic analysis of the relationships of selected vespertilionine species (chiefly those currently 
referred to Pipistrellus and Eptesicus) by Horacek & Hanak (1985-1986) appeared while this paper 
was in press. These authors provided definitions of Pipistrellus, Hypsugo (which they considered 
generically valid) and Eptesicus, based on the morphology of the penis and baculum, the upper 
molars, the basisphenoid pits, the pelvic girdle, and the tibia, tail and epiblema. 

Horacek & Hanak suggested that the classification of pipistrelloid bats might be clarified by the 
recognition of additional subgenera or genera for those species or species groups that do not 
conform precisely with those that they included within these three generic groupings. To some 
extent such recognition is provided in several instances by the classification here proposed and 
although some major differences exist between the informal assessments and species groups of 
Horacek & Hanak and the formal arrangement put forward in this paper there is nevertheless 
a broad measure of agreement. Horacek & Hanak did not attempt any classification of the 
Vespertilioninae as a whole, but 'Nycticeius' schlieffenii, here considered to represent a distinct 
monospecific genus (Nycticeinops gen. nov.) was thought by these authors to be referable either to 
Eptesicus ( Rhyneptesicus), or possibly to justify the establishment of a new subgenus within 
Eptesicus. 



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Academy of Scientific Research and Technology Natural History Museum of Egypt. 
Wettstein, O. van 1916. Neue Affen und Fledermause aus Nordost-Afrika. Anzeiger der f Kaiser lichen J 

Akademie der Wissenschaften. Mathematische-Naturwissenschaftliche Klasse, Wien 53: 189-192. 
Williams, D. F. & Mares, M. A. 1978. Karyologic affinities of the South American big-eared bat, Histiotus 

montanus (Chiroptera: Vespertilionidae). Journal of Mammalogy, Baltimore 59: 844-846, 1 fig. 
Wimsatt, W. A. & K alien, F. C. 1952. Anatomy and histophysiology of the penis of a vespertilionid bat, 

Myotis lucifugus lucifugus, with particular reference to its vascular organisation. Journal of Morphology, 

Boston 90: 41 5-465, 4 figs, 6 pis. 
Zima, J. & Horaeek, I. 1985. Synopsis of karyotypes of vespertilionid bats (Mammalia: Chiroptera). Acta 

Universitatis Carolinae-Biologica, Pragae (1981): 31 1-329, 1 fig., 2 tabs. 
Zubaid, A. & Davison, G. W. H. (In press). A comparative study of the baculum in Peninsular Malaysian 

hipposiderines. Mammalia, Paris. 

Manuscript accepted for publication 30 June 1986. 



VESPERTILIONINE SYSTEMATICS 269 

Appendix 1. Specimens examined 

AMNH = American Museum of Natural History, New York 
BM(NH) = British Museum (Natural History), London 
CMNH = Carnegie Museum of Natural History, Pittsburgh 
HZM = Harrison Zoological Museum, Sevenoaks, Kent 
NMW = Naturhistorisches Museum, Wien 

Vespertilioninae 

Myotis nattereri 

HZM 26.1 1254 Leany Cave, Pilis Heights, Hungary. (Fig. 19j) 

Myotis ridleyi 

BM(NH) 98.3.13.5 Selangor, Malaya. Holotype. (Fig. 19i) 

Pizonyx vivesi 

HZM 3.10284 Isla Cordonosa, Bahia de Los Angeles, Baja Norte, Mexico. (Fig. 19k) 

Lasionycteris noctivagans 

BM(NH) 7.7.7.2319 Raleigh, North Carolina, USA. (Fig. 170 

HZM 2.3708 Delta, Manitoba, Canada 

Plecotus auritus 

HZM 19.1227 Near Godstone, Surrey, England. (Fig. 19g) 

Plecotus austriacus 

BM(NH) 91.10.5.4 Duirat, Tunis 

HZM 3.4867 St. Pierre de Varenne, Saone et Loire, France. 

HZM 4.8337 Mont de Lans, Les Deux Alpes, Isere, France. 

HZM 5.8467 Chateau de Salse, Salse, Rousillon, France. (Fig. 19h) 

Barbastella barbastellus 

HZM 13.1 1222 Kiralyret, Borzsony Heights, Hungary. (Fig. 18j) 

Rhogeessa tumida 

HZM 1.12080 Airport Camp, Belize. (Fig. 18k) 

Nycticeius humeralis 

AMNH 249144 Sierra de Tamaulipas, Acuna, Tamaulipas, Mexico, 2890 ft. (Fig. 17k) 

Otonycteris hemprichii 

BM(NH) 14.8.17.1 Syrian Desert. 

HZM 6.8174 17 km N of Hufoof, Saudi Arabia. (Fig. 16a) 

Lasiurus cinereus 

HZM 1.3695 S Fork, Cave Creek, near Portal, Cochise County, Arizona, USA. (Fig. 191) 

Dasypterus argentinus 

BM(NH) 33.6.24.3 Bonifacio, Argentina. (Fig. 180 

Antrozous pallidus 

BM(NH) 50.767 California, USA. (Fig. 18b). 

HZM 3.3692 S Fork, Cave Creek, near Portal, Cochise County, Arizona, USA. 

Scotophilus borbonicus 

BM(NH) 89.1.11.2 E coast of Africa. 

Scotophilus dinganii 

BM(NH) 79.513 Tokadeh, Nimba, Liberia, 600 m. 

Scotophilus heathii 

BM(NH) 14.7.19.19 Mount Popa, Burma. 

BM(NH) 14.7.19.28 Kyauk Inyaung, Irrawaddy, Burma. 

BM(NH) 60.257 Tori, Pakistan. (Fig.lTh) 

BM(NH) 70.1488 Bang Phra, Cholburi, Sriracha, Thailand. 

BM(NH) 76.787-788 Nhatrang, Annam, Vietnam. 



270 J. E. HILL & D. L. HARRISON 

Scotophilus kuhlii 

BM(NH) 75.2955 Chiang Mai, Thailand (Fig. 17i) 

Scotophilus nigrita (gigas) 

BM(NH) 22.12.17.55a Mtondo, Ruo, Malawi. (Fig. 17g) 

Scotophilus nigritellus 

BM(NH) 78.189 Numan, Gongola, Nigeria. (Fig. 17j) 

Scotomanes ornatus 

BM(NH) 94.9. 1.21 Foochow, China. (Fig. 18g) 

Eptesicus (Eptesicus) 

Eptesicus bobrinskoi 

BM(NH) 63.1 187 Outer Su, N of Mount Sabalan, NW Iran. (Fig. 13e) 

Eptesicus nasutus 

HZM 3.4571 Harmul, 10 m N of Sohar, Oman (batinensis). 
HZM 12. 1 1 172 Jamma, near Rostaq, Oman (batinensis). (Fig. 14c) 
HZM 1.1623 Shaiaba, Iraq (pellucens). 

Eptesicus bottae 

HZM 18.1616 Ser'Amadia, Kurdistan, Iraq. 

HZM 5.1628 Basrah, Iraq (hingstoni). 

BM(NH) 3.12.8.9 Cairo, Egypt (inncsi). (Fig. 13j) 

HZM 12.8075 Birkat Sharaf al Wadi Sahtan, Jebel al Akhdar, Oman (omanensis). (Fig. 14a) 

Eptesicus brasiliensis 

BM(NH) 85.6.26.10 San Lorenzo, Rio Grande do Sul, Brazil. (Fig. 13k) 

BM(NH) 0.6.29.4 Palmeira, Parana, Brazil. 

BM(NH) 98.10.3.32 Valdivia, Colombia (andinus). (Fig. 13d) 

Eptesicus furinalis 

BM(NH) 4.8.8.5 La Plata, Argentina. (Fig. 13c) 

Eptesicus fuscus 

BM(NH) 89.6.1.4 Sing Sing, New York, USA. (Fig. 13a) 

BM(NH) 52.551 Chinchona, Jamaica (hispaniolae). (Fig. 13i) 

Eptesicus hottentotus 

BM(NH) 81.7.1 1.1 Drakenburg Mountains, Natal, South Africa (megalurus). (Fig. 13b) 

Eptesicus serotinus 

BM(NH) 53.555 Blandford, Dorset, England. (Fig. 13g) 

HZM 3.629 Shepreth, Cambridgeshire, England. 

BM(NH) 66.1 150 Defilia Oasis, Figuig, Morocco (isabellinus}. (Fig. 13h) 

Eptesicus (Rhinopterus) 

Eptesicus floweri 

BM(NH) 0.8.6.20 Abu Zeit, White Nile, Sudan. (Fig. 130 

BM(NH) 1.5.5.78 Shendy, Sudan (lowei). (Fig. 131) 

Vespertilio orientalis 

BM(NH) 8.7.25.6, BM(NH) 8.8.1 1.2 Kuatun, NW Fokien, China 

Histiotus macrotis 

BM(NH) 71.1 123 Antofagasta, Lake Miniques, Chile, 1450 m. (Fig. 18e) 

Histiotus (?) macrotis 

BM(NH) 6.5.8.3 Jafi, Tucuman Province, Argentina. (Fig. 18d) 

Histiotus velatus 

BM(NH) 0.6.29.2 Palmeira, Parana, Brazil. (Fig. 18c) 

Tylonycteris pachypus 

BM(NH) 9.1. 5.954 Buitenzorg, Java. (Fig. 18h) 



VESPERTILIONINE SYSTEMATICS 271 

Tylonycteris robustula 

BM(NH) 60.1499 Bukit Lagong Forest Reserve, Kepong, Selangor, Malaya. (Fig. 18i) 

HZM 3.7444 15th mile Ulu Gombok, Selangor, Malaya. 

Mimetillus moloneyi 

BM(NH) 93. 1 .7.2 Leekie, Nigeria 

BM(NH) 54.862 Irumu, Zaire. 

BM(NH) 60.154 Bo, Sierra Leone. 

BM(NH) 64.1788 Liwale, Tanzania. 

HZM 2.7802 Near Babeke, River Isai, Ituri, Zaire. 

Glauconycteris argentata 

BM(NH) 54.863 Banana, Zaire. 

BM(NH) 59.510 Ikela, Ikela Territory, Zaire. (Fig. 19d) 

Glauconycteris beatrix 

BM(NH) 48.713 Eshobe, Mamfe, Cameroun. (Fig. 19c) 

Glauconycteris humeralis 

BM(NH) 30.11.11.173 River Wasa, Semliki Valley, Uganda. (Fig. 1 9e) 

Glauconycteris poensis 

BM(NH) 96. 1 2.3 1 .2 Sierra Leone. 

BM(NH) 69.26 Abidjan, Ivory Coast. (Fig. 19a) 

Glauconycteris variegata 

BM(NH) 76.780 Mole National Park, Ghana. (Fig. 19b) 

BM(NH) 55.409 Mongue, near Inhambane, Mozambique (papilio). (Fig. 190 

Pipistrellus ( Pipistrellus ) 

Pipistrellus pipistrellus 

HZM 94.6807 Rabat, Malta. 

HZM 1 16.8549 Sevenoaks, Kent, England. (Fig. 2a) 

HZM 1 17.8650 Aylesford, Kent, England. 

BM(NH) 73.689 Kululai Rest House, Northwest Frontier Province, Pakistan (bactrianus) 

BM(NH) 14.5.10.19 BM(NH) 14.5.10.22 Djarkent, Semiretschenskoi, USSR (lacteus). 

Pipistrellus nathusii 

BM(NH) 8.8.4.128 BM(NH) 62.1368 St Giles, France (Fig. 2b) 

Pipistrellus abramus 

BM(NH) 89.6.17.3-4 Kin Kiang, Yangtse Kiang, China (seen by Thomas, 1928a). 

BM(NH) 5. 1 .4.8 Tokyo, Japan. 

BM(NH) 7.7.3.26 Nanking, China (seen by Thomas, 19280). (Fig. 3a) 

BM(NH) 14.10.1.1 Lokow, Hunan, C China. 

BM(NH) 26. 1 0.4. 1 8 Hue, Annam, Vietnam (seen by Thomas, 1 928a) 

BM(NH) 66.3469-3470 Chihli, China. 

BM(NH) 86.529 Chusan, China (Syntype irretitus). 

BM(NH) 86.532 Canton, S China. 

Pipistrellus babu 

BM(NH) 45. 1.8.403 Nepal. 

BM(NH) 16.3.25.8 Pashok, Darjeeling, India. (Fig. 4a) 

Pipistrellus camortae 

BM(NH) - - Car Nicobar (Original No. 3/76). (Fig. 1 5d) 

Pipistrellus endoi 

BM(NH) 70.2522 Horobe, Tayama, Ajiro-Machi, Minohe-Gun, Iwate Prefecture, Japan. (Fig. 3b) 

Pipistrellus javanicus (tralatitius) 

BM(NH) 0.8.2.9 Sumatra. 

BM(NH) 9. 1 .5.295 Tjilatjap, Java. 

BM(NH) 9. 1 .5.997-998 W Java. 

BM(NH) 16.4.21.3 Sungei Penoh, Korinchi, Sumatra. 



272 J. E. HILL & D. L. HARRISON 

BM(NH) 27.12.1.37 Tarn Dao, Tonkin, Vietnam, 3000 ft (No. 41 1 of Thomas, 1928a). 

BM(NH) 28.7. 1 .20 Phu Qui, Annam, Vietnam, 100 ft (No. 866 of Thomas, 19286, who identified the specimen 

as P. coromandra tramatus, but with a longer baculum than those previously examined). 
BM(NH) 83.76 Silau Silau Trail, Mount Kinabalu, Sabah, Borneo. (Fig. lOe) 

Pipistrellus paterculus 

BM(NH) 14.7.8.62 Pyaunggaung, N Shan States, Burma, 2794 ft. 
BM(NH) 14.7.19.241 Kyauk Myaung, Irrawaddy, W Burma. 
BM(NH) 14.7.19.242 Mount Popa, Upper Burma (Holotype). (Fig. 3c) 
BM(NH) 14.7.19.240 Mandalay, Burma. 

Pipistrellus angulatus 

BM(NH) 67.2125 Schoolmaster's House, Nuhu, Guadalcanal I, Solomon Is (ponceleti). (Fig. 4d) 

Pipistrellus collinus 

BM(NH) 50.983 Baiyanka, Purari-Ramu Divide, SE Bismarck Range, Papua New Guinea. (Fig. 4b) 

Pipistrellus coromandra 

BM(NH) 32. 1 1 . 1 .7 Nam Tamai, Upper Burma. 

BM(NH) 50.478 Ningma, Upper Burma. 

BM(NH) 76.1263 Sumka Uma, Upper Burma. 

HZM 1.7317, HZM 2.73 18 Near Mirzapur, India. (Fig. 7c, HZM 2.7318) 

HZM 4.7320 Dalatpur, near Mirzapur, India. 

BM(NH) 4.6.8.1 Annam, Vietnam (tramatus). (Fig. 7b) 

BM(NH) 27.12.1.40 Bac-kan, Tonkin, Vietnam (tramatus) (Original No. 444, seen by Thomas, 1928a). 

Pipistrellus mimus 

BM(NH) 98.5.5.20 Dangs, Bombay, India. 

HZM 1 .10456 Vikas Vidyalaya, near Ranchi, Bihar, India. (Fig. 7g) 

Pipistrellus murrayi 

BM(NH) 99.8.6.34 Christmas I, Indian Ocean (Holotype). (Fig. 4c) 

BM(NH) 9.1.16.7 Flying Fish Cove, Christmas I, Indian Ocean. 

Pipistrellus papuanus 

BM(NH) 22.2.2.3 Fredrik Hendrik I, Irian Jaya. (Fig. 2c) 

BM(NH) 34.1.14.8 Kokoda, Papua New Guinea. 

Pipistrellus tenuis 

BM(NH) 85.912 Coast of Sabah, Borneo (nitidus). (Fig. 9d) 

Pipistrellus ceylonicus 

BM(NH) 95.6.12.1 Pundibiya, India. 

BM(NH) 2.4.2.8 Astoli, Belgoum, India. (Fig. 7d) 

BM(NH) 1 1.4.5.5 Lanje, Konkan, India. 

BM(NH) 13.9.8.102 Gujerat, India. 

BM(NH) 9.1.4.73 Mangalore, Malabar Coast, India (Holotype indicus) 

BM(NH) 4.6.8.7-8 Tonkin, Vietnam (raptor). (Fig. 3d, BM(NH) 4.6.8.7 Holotype). 

Pipistrellus crassulus 

BM(NH) 4.2.8.1 Efulen, Cameroun (Holotype). (Fig. 7e) 

Pipistrellus nanulus 

BM(NH) 4.2.8.8 Efulen, Cameroun (Holotype). (Fig. If) 

BM(NH) 79.508 South Nimba, Liberia. 

Pipistrellus rueppellii 

BM(NH) 68.12.22.3 Zanzibar (Holotype pulcher). (Fig. lOa) 

BM(NH) 99.9.9.20 Egypt. 

BM(NH) - - Uganda. (Fig. lOb) 

HZM 3.3170 Kabompo Boma, Zambia. 

HZM 7. 12109 Suez, Egypt. 

Pipistrellus deserti 

BM(NH) 79.987 Hoggar Plateau, Algeria. (Fig. 5c) 

NMW 27503 (?) Upper Egypt. 



VESPERTILIONINE SYSTEMATICS 273 

Pipistrellus kuhlii 

BM(NH) 92.9.9.25 Upper Egypt. 

BM(NH) Argostoli, Cephaloni, Greece. (Fig. 5a) 

BM(NH) 63.335 Sangha, Malya Khola, E Nepal. 

HZM 5. 1 1607 Horefto, near Volos, Greece. 

HZM 11.1016 Rapallo, N Italy. 

HZM 138.4563 Yal bu Hillal, Batinah, Oman. 

HZM 154.4619 Saham, Batinah, Oman. 

HZM 203.7232 Dig Dagga, Ras al Khaima, United Arab Republic. 

HZM 218.7402 Benghazi, Libya. 

HZM 227.91 10 Kapsowat, Marakwat, Kenya. 

Pipistrellus maderensis 

BM(NH) 86.528 Madeira. (Fig. 5b) 

Pipistrellus rusticus 

BM(NH) 35.9.1.108 Okavango-Omatako Junction, Grootfontein District, Namibia. 

BM(NH) 79.1731 Oli River, Borgu G.R., Nigeria. (Fig. 6c) 

HZM 4.3285 Sentinel Ranch, River Limpopo, Zimbabwe. (Fig. 5d) 

Pipistrellus ( Vespadelus} 

Pipistrellus pumilus 

BM(NH) 70.1093 E Bonithon Range, C Australia 2342'S, 12902'E, 1400 ft. 

BM(NH) 71.1497 Westwood, near Rockhampton, Queensland, Australia. (Fig. 12k) 

Pipistrellus (Perimyotis) 

Pipistrellus subflavus 

HZM 1.2422 Big Wyandotte Cave, Crawford County, Indiana, USA. (Fig. 2d) 

Pipistrellus (Hypsugo) 

Pipistrellus anchietae 

BM(NH) 69.1248 Ngoma, Zambia. 

BM(NH) 70.2632 Balovale, Zambia. (Fig. 6e) 

BM(NH) 89.5.1.5 Caconda, Angola (Syntype of Vesperus bicolor Bocage, 1889). (Fig. 9e) 

Pipistrellus bodenheimeri 

HZM 3.3786 Jazirat al Abid, Aden, South Yemen. 

HZM 5.8279 Bin Gedi, Israel. (Fig. 90 

Pipistrellus savii 

BM(NH) 31.11.11.13, BM(NH) 66.4644 E slope of Mount Olympus, Greece. 

BM(NH) 61.395 Ainab, Lebanon. (Fig. 6a) 

Pipistrellus arabicus 

HZM 4.10060 Wadi Sahtan, Oman. 

HZM 5.1 1625 Wadi Fidah, Dank/Ibri, Oman. (Fig. 7a) 

Pipistrellus helios 

BM(NH) 39.133 N Guaso Nyiro, Kenya. (Fig. 6d) 

BM(NH) 69.207 Kangatet, S Turkana, Kenya. 

HZM 2.4086 Archer's Post, Northern Frontier District, Kenya. 

Pipistrellus nanus 

BM(NH) 49.484 Kontaur, Gambia. 

HZM 3.2778 Sokoto, N Nigeria. 

HZM 3.4026, HZM 4.4027 Near Monrovia, Liberia. 

HZM 83.4387 Haroni-Lusitu Beacon 74, Zimbabwe. 

HZM 107.3212 Kabompo Boma, Zambia. 

HZM 146.5161, HZM 147.5162 Rondo, Lindi, Tanzania. 

HZM 165.5321 Liwale, Tanzania. 

HZM 200.6581 Karonga, Malawi. 

HZM 258.1 1469 Kunyale Stream, Mwinilunga District, Zambia. 



274 J. E. HILL & D. L. HARRISON 

HZM 260.12175 Lamto, Ivory Coast. 

HZM 26 1 . 1 2 1 76 Ivory Coast. 

HZM 263.12451, HZM 264.12452 Kamuani Area, Machakos District, Kenya. (Fig. 6b, HZM 263.12451) 

Pipistrellus pulveratus 

BM(NH) 79.702 Near Nicholson Goat Bungalows, Hong Kong I. 

BM(NH) 79.903 Peace Mansion, Tai Hang Road, New Territories, Hong Kong (Fig. 8c) 

Pipistrellus Hesperus 

BM(NH) 98.3.1.8 Sierra Laguna, Baja California, Mexico. 

BM(NH) 29.1 1.7.10 Panamint Mts, California, USA. 

HZM 4.1 1219 Sycamore Well, Hidalgo County, New Mexico, USA. (Fig. 8d) 

Pipistrellus eisentrauti 

BM(NH) 84.1684, BM(NH) 84.1686 Mount Cameroun, Cameroun. (Fig. 9g, BM(NH) 84.1684) 

Pipistrellus imbricatus 

BM(NH) 9.1.5.286 Buitenzorg, Java. (Fig. 9a) 

Pipistrellus macrotis 

BM(NH) 23.1.2.12 Sabang, NW Sumatra. (Fig. 9b) 

Pipistrellus kitcheneri 

BM(NH) 10.4.5.47 Boentok, Barito River, Kalimantan, SC Borneo. (Fig. 8e) 

Pipistrellus lophurus 

BM(NH) 14.12.1.6 Maliwun, Victoria Province, Tenasserim, Burma (Holotype). (Fig. 8f) 

Pipistrellus stenopterus 

BM(NH) 60.1537 Institute of Medical Research Compound, Kuala Lumpur, Malaya. 

BM(NH) 65.135 Pasir Road, Kuala Lumpur, Malaya. (Fig. 7h) 

Pipistrellus (Falsistrellus) 

Pipistrellus affinis 

BM(NH) 83.3.3.2 Wynaard, India. (Fig. 8a) 

BM(NH) 72.4224 Argarawa, Nevrawa Elwa, Central Province, Sri Lanka. 

Pipistrellus peter si 

BM(NH) 23. 1 .2.3. Buru I, Molucca Is (Fig. 8b) 

Pipistrellus tasmaniensis 

HZM 1.8712 Barrington Tops National Park, New South Wales, Australia. (Fig. 8g) 

Pipistrellus (Neoromicid) 

Pipistrellus capensis 

BM(NH) 32.9.1.249 Broken Hill, Zambia. 

BM(NH) 54.859 Elizabethville, Zaire. 

BM(NH) 61.1078 Doddieburn Ranch, West Nicholson, Zimbabwe, 2300 ft, 2124'S, 2921'E. 

BM(NH) 72.4383 E of Lake Margharita, Bulcha Forest, Ethiopia, 1800 m, 06 1 1'N, 3610'E. 

BM(NH) 72.4391 Didessa River, Wollega Province, Ethiopia, 1 190 m, 0902'N, 3609'E. (Fig. 12g) 

BM(NH) 75.561 Mole National Park, Ghana. (Fig. 12b) 

BM(NH) 83.200 Mcheni Gorge, Chizarira National Park, Binga Province, Zimbabwe, 1740'S, 2752'E. 

HZM 36.4514 40 m NW of Serowe, Botswana. 

BM(NH) 66.6057 Ambositra, Madagascar (matrokd). (Fig. 12a) 

BM(NH) 77.2.19.6 Anzahameru, Madagascar ('minutus'). (Fig. 12i) 

Pipistrellus guineensis 

BM(NH) 70.2224, BM(NH) 70.2228, BM(NH) 72.4373 Gambela, Ethiopia, 815'N, 3435'E (BM(NH) 

72.4373 at 515 m) (Fig. 12c, BM(NH) 70.2224) 
BM(NH) 76.293 Shagamu, Nigeria. 
BM(NH) 84.1019 Bontioli, Bougouriba River, Burkina Faso (Upper Volta). 

Pipistrellus melckorum 

BM(NH) 83.216 Mcheni Gorge, Chizarira National Park, Binga Province, Zimbabwe, 1740'S, 2752'E. 
(Fig. 12Q 



VESPERTILIONINE SYSTEMATICS 275 

Pipistrellus somalicus 

BM(NH) 70.484 Mouth of Fincha River, Blue Nile Gorge, Ethiopia, 1003'N, 3720'E. (Fig. 12h) 

BM(NH) 76.814 S bank of Ganale Doria, Sidam-Bale Bridge, Sidamo Province, Ethiopia, 545'N, 3937'E. 

BM(NH) 84.1016 Comoe River, Burkina Faso (Upper Volta), 260 m, 957'N, 438'W. 

CMNH MJS 2846 Snai Sugar Plantation, l km S, \ km E of Giohar, Somalia, 246'N, 453 1 'E. 

Pipistrellus zuluensis 

BM(NH) 83.212 Mchesu River, Chizarira National Park, Binga Province, Zimbabwe, 1747'S, 2739'E. 

BM(NH) 83.215 Singama, Sibuwa, Binga Province, Zimbabwe, 17 C 36'S, 2751'E. (Fig. 12d) 

Pipistrellus rendalli 

BM(NH)89.12.12.1 Bathurst, Gambia. 

BM(NH) 7.12.17.1-2 Gondokoro, White Nile, Sudan. 

BM(NH) 23.4.12.1-2 Bugala, Sesse Is, Victoria Nyanza, Uganda. (Fig. 12e, BM(NH) 23.4.12.2) 

BM(NH) 48.702 N'ko, Obubra Division, S Nigeria (Ibrunneus). (Fig. 14b) 

Pipistrellus tenuipinnis 

BM(NH) 47.350 Umuahia, E Nigeria. 

BM(NH) 54.917 Bonthe, Sierra Leone. 

BM(NH) 67.1734 Bota, Victoria, Cameroun, 400'N, 905'E. (Fig. 12j) 

Pipistrellus (Arielulus) 

Pipistrellus circumdatus 

BM(NH) 73.618 Telecommunications Tower, Fraser's Hill, Pahang, Malaya. (Fig. 2e) 

Pipistrellus cuprosus 

BM(NH) 83.351 Sepilok, Sabah, Borneo, 552'N, 1 1756'E (Holotype). (Fig. 9h) 

Pipistrellus societatis 

BM(NH) 67.1605 Base Camp, Gunong Benom, Pahang, Malaya, 800 ft (Holotype). (Fig. 9c) 

Nyctalus noctula 

BM(NH) - - Locality unknown. 

HZM 10.613 Bottisham, Cambridgeshire, England. 

HZM 33.8888 Winchelsea Beach, Sussex, England. (Fig. 100 

Laephotis botswanae 

BM(NH) - - Zomba, Malawi (original No. 2269; damaged). 

Laephotis wintoni 

HZM 1.3020 Nyeri, Mount Kenya, Kenya. (Fig. 160 

Glischropus tylopus 

BM(NH) 10.4.5.136 Upper Barito River, Kalimantan, SC Borneo. (Fig. 18a) 

Scotozous dormeri 

BM(NH) 12.3.8.30 Furdapur, Ajanta, Khandesh, India. 

BM(NH) - - Kathiawar, India (Original No. BNHS 2007). (Fig. 1 6d) 

Scoteanax rueppellii 

BM(NH) 80.3.25.1 Richmond River, New South Wales, Australia. (Fig. 16i) 

Scotorepens balstoni 

BM(NH) 10.6.21.9 Hermannsburg, Northern Territory, Australia. (Fig. 16g) 

Scotorepens greyii 

BM(NH) 75.2261 Pine Creek, 20 m ESE of Candy's Hill, Northern Territory, Australia, 1349'S, 13149'E. 
(Fig. 16h) 

Nycticeinops schlieffenii 

BM(NH) 14.7.3 1.14 Wei Wei River, Kenya. 

BM(NH) 15.3.6.66 Kamisu, Dinda River, Sudan. 

BM(NH) 71.675 Awash, Filhoa, Ethiopia, 0900'N, 3858'E. 

HZM 5.2120 Ikau, Rukwa, Tanzania. (Fig. 16e) 

Scotoecus albigula 

BM(NH) 63.1042 Calundo, Lunda, Angola. (Fig. 20a) 



276 J. E. HILL & D. L. HARRISON 

Scotoecus albofuscus 

BM(NH) 96. 1 2.3 1 . 1 Sierra Leone. (Fig. 20e) 

Scotoecus hindei 

BM(NH) 14.7.31.13 30m NW of Baringo, Kenya. (Fig. 20d) 

BM(NH) 66.1466 Jos, Nigeria (falabae). (Fig. 20b) 

Scotoecus hirundo 

BM(NH) 76.771 Mole National Park, Ghana. (Fig. 20c) 

Scotoecus pallidus 

BM(NH) 86.531 Afghanistan (damaged). 

Philetor brachypterus 

BM(NH) - - New Guinea. (Fig. 16b) 

Hesperoptenus (Milithronycteris) 

Hesperoptenus blanfordi 

BM(NH) 83.853 Sepilok, Sabah, Borneo, 552'N, 1 1756'E. (Fig. 21g) 

Hesperoptenus tickelli 

BM(NH) 71.12.26.1 Sri Lanka. (Fig. 21b) 

Hesperoptenus tomesi 

BM(NH) 7.1.1.428 Malacca, Malaya (Holotype). (Fig. 2 la) 

Chalinolobus gouldi 

BM(NH) 71.1504 Westwood, near Rockhampton, Queensland, Australia. (Fig. 17b) 

Chalinolobus morio 

BM(NH) 6.8.1.60 (King River, Western Australia. (Fig. 17a) 

Chalinolobus nigrogriseus 

BM(NH) 44.6.13.2 Port Essington, Northern Territory, Australia (rogersi). 

BM(NH) 75.2260 Pine Creek, 20 m ESE of Gandy's Hill, Northern Territory, Australia, 1349'S, 13149'E. 
(rogersi). (Fig. 17c) 

Chalinolobus picatus 

BM(NH) 9.3.7.2 Gunnamulla, Queensland, Australia. (Fig. 17d) 

Chalinolobus tuberculatus 

BM(NH) 89.10.27.1 Outlying islands near Stewart I, New Zealand. (Fig. 17e) 

Nyctophilinae 

Nyctophilus bifax 

BM(NH) 67.5.6.5 Cape York, Queensland, Australia. 

BM(NH) 77.3.28.1 Islands of Torres Straits, Australia. 

BM(NH) 86.1 1.8.12 Somerset, Cape York, Queensland, Australia. 

BM(NH) 15.3.13.1 Cloncurry, Queensland, Australia. 

BM(NH) 15.3.13.3 Herberton District, Queensland, Australia (Holotype). (Fig. 22a) 

Nyctophilus daedalus 

BM(NH) 47.7.2 1 . 1 6, BM(NH) - - Port Essington, Northern Territory, Australia. (Fig. 22g, BM(NH) 

47.7.21.16) 
BM(NH) 97.4.12.5 Daly River, Northern Territory, Australia. 

Nyctophilus gouldi 

BM(NH) 15.3.13.7 Ash I, Hunter River, New South Wales, Australia (damaged, part lost). 

BM(NH) 15.3.13.8 Sydney, New South Wales, Australia. 

BM(NH) - - Botany, Sydney, New South Wales, Australia (Original No. 164) (Fig. 22d) 

HZM 1.12085 Werrikimbe, Hastingsshire, New South Wales, Australia. (Fig. 16c) 

BM(NH) 52. 1 . 1 5.30 Tasmania (sherrini). (Fig. 220 

Nyctophilus geoffroyi 

BM(NH) 15.3.13.1 1 Kosciusko, New South Wales, Australia (pacificus). 

BM(NH) - - Tasmania (pacificus) (Original No. M.I 735). 



VESPERTILIONINE SYSTEMATICS 277 

BM(NH) - - Launceston, Tasmania (pacificus) (Original No. M. 168) (Fig. 22e) 

BM(NH) 7.1.4.3 Alexandria, Northern Territory, Australia (pallescens). (Fig. 22b) 

Nyctophilus microtis 

BM(NH) 88.4.18.1 Sogeri, Papua New Guinea (Holotype). (Fig. 22c) 

Pharotis imogene 

BM(NH) 97.8.7.21 Kamali, Papua New Guinea. (Fig. 22h) 



278 



J. E. HILL & D. L. HARRISON 





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VESPERTILIONINE SYSTEMATICS 



279 



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280 



J. E. HILL & D. L. HARRISON 



Table 2 Usual incisive and premolar dental formulae in the Vespertilioninae and Nyctophilinae. Total 
number of teeth (including four canines and twelve molars) in parentheses. Dental notation of Miller ( 1 907). 



(38) Myotis, Pizonyx 

(36) Lasionycteris, Plecotus, Idionycteris, Eudiscopus 

(34) Euderma, Barbastella, la, Pipistrellus, Glischropus, Scotozous, Nyctalus, 

Chalinolobus 
(32) Eptesicus, Vespertilio, Histiotus, Tylonycteris , Mimetillus, Glauconycteris, 

Pipistrellus, Laephotis, Philetor, Hesperoptenus 
(32) Lasiurus 
(30) Rhogeessa, Baeodon, Nycticeius, Otonycteris, Dasypterus, Scotomanes, 

Scotophilus, Scoteanax, Scotorepens, Nycticeinops, Scotoecus, 

Nyctophilus, Pharotis 
(28) Antrozous, Bauerus 



Table 3 Classifications of the Vespertilioninae and Nyctophilinae. That of Tate (\942a) is concerned primarily with 
Oriental and Australasian taxa, those of Koopman with Australasian (1973) and predominantly African (1975) forms. 



Tate(1942a) 



Koopman (1973, 1975) 



Hill & Harrison 



Pipistrellus 
abramus group 
abramus 
akokomuli 
bancanus 
camortae 
irretitus 
paterculus 
pumiloides 
pipistrellus group 

pipistrellus (Including 

bactrianus) 
nathusii 

coromandra group 
aladdin 
angulatus 
collinus 
coromandra 
imbricatus 
meyeni 
micropus 
murrayi 
ponceleti 
portensis 
regulus 
sturdeei 
subulidens 
tramatus 
tenuis group 
mimus (Including 

glaucillus) 
nitidus 
papuanus (Including 

orientalis) 
principulus 
tenuis 
ceylonicus group 

ceylonicus (Including 

chrysothrix, indicus, 

subcanus) 



Pipistrellus 

Amalgamates pipistrellus, abramus 
( =javanicus), coromandra and 
tenuis groups of Tate (1942a) 
pipistrellus group 
imbricatus 
javanicus (Including 

abramus) 
meyeni 
nanus (Including 

(?) Helios) 
permixtus 

tenuis (Including angulatus, 
collinus, nitidus, papuanus, 
ponceleti, murrayi, sewelanus, 
subulidens, westralis [Koopman, 
1984c]) 



ceylonicus group 
ceylonicus 



Pipistrellus 

Pipistrellus (Pipistrellus) 
pipistrellus group 
pipistrellus subgroup 
pipistrellus (Including aladdin, 

bactrianus, lacteus, 

mediterraneus) 
nathusii 
permixtus 
javanicus subgroup 
abramus (Including 

akokomuli, irretitus, 

pumiloides) 
babu 
endoi 
javanicus (Including bancanus, 

camortae, meyeni, 

'tralatitius") 
paterculus 
peguensis 

coromandra subgroup 
adamsi 

angulatus (Including ponceleti) 
collinus 
coromandra (Including afghanus, 

portensis, tramatus) 
mimus (Including 

glaucillus, principulus) 
murrayi 
papuanus 
sturdeei 
tenuis (Including nitidus, 

sewelanus, subulidens) 
wattsi 
westralis 

ceylonicus subgroup 
ceylonicus (Including borneanus, 

chrysothrix, indicus, raptor, 

shanorum, subcanus) 
(?) minahassae 



VESPERTILIONINE SYSTEMATICS 



281 



Table 3-cont. 



Tate(1942a) 



Koopman (1973, 1975) 



Hill & Harrison 



minahassae group 
minahassae 

rueppellii group 
coxi 



kuhlii group 
babu 
canus 
kuhlii (Including 

ikhwanius, lepidus) 
leucotis 
lobatus 



Eptesicus 
pumilus group 
pumilus (Including 
caurinus, darlingtoni, 
vulturnus) 
pygmaeus 



Pipistrellus 
savii group 
austenianus 
cadornae 
curtatus 
macrotis 
savii 
vordermanni 



minahassae group 

minahassae 
rueppellii group 

rueppellii (Including (?) 
fuscipes; pulcher) 



kuhlii group 
aero 

anchietae 
deserti 
inexspectatus 
kuhlii (Including (?) 

aegyptius; fuscatus) 
rusticus (Including marrensis) 



savii group 
ariel 
macrotis 
maderensis 



Hesperus group 
hesperus 
musciculus 



joffrei group 
anthonyi 
joffrei 
stenopterus 



joffrei group 
stenopterus 



rueppellii group 
crassulus 
nanulus 

rueppellii (Including coxi, 
fuscipes, leucomelas, pulcher, 
senegalensis , vernayi) 
kuhlii group 
aero 
deserti 

inexspectatus 
kuhlii (Including (?) 
aegyptius; fuscatus, 
ikhwanius) 
maderensis 

rusticus (Including marrensis) 
Pipistrellus ( Vespadelus) 
douglasorum 
pumilus (Including 

darlingtoni) 
regulus 
sagittula 
vulturnus 
Pipistrellus (Perimyotis) 

subflavus 

Pipistrellus (Hypsugo) 
savii group 
savii subgroup 
anchietae ( = 'bicolor'7) 
ariel 

austenianus 
bodenheimeri 
savii (Including caucasicus, 

darwini, maurus) 
nanus subgroup 
arabicus 
helios 
musciculus 
nanus (Including culex, 

stampflii) 
pulveratus subgroup 

pulveratus 
hesperus subgroup 

hesperus 
eisentrauti subgroup 

eisentrauti 
imbricatus subgroup 
curtatus 
imbricatus 
macrotis 
vordermanni 
lophurus subgroup 
cadornae 
kitcheneri 
lophurus 

stenopterus group 
anthonyi 
joffrei 
stenopterus 



282 

Table 3-c ont. 



J. E. HILL & D. L. HARRISON 



Tate(1942fl) 



Koopman( 1973, 1975) 



Hill & Harrison 



affinis group 


affinis group 


Pipistrellus (Falsistrellus) 


affinis 


kitcheneri 


affinis group 


kitcheneri 


petersi 


affinis 


lophurus 




(?) mordax 


petersi 




petersi 


pulveratus 






tasmaniensis group 




tasmaniersis group 


tasmaniensis (Including 




mackenziei 


krefftii) 




tasmaniensis (Including 



circumdatus group 

circumdalus 

mordax 
Eptesicus 

Eptesicus (Amblyotus) 

alaschanicus 

bobrinskoi 



matschiei (Including 

pellucens) 
nilssonii (Including 

caucasicus, centrasiaticus, 
gobiensis, kashgaricus, 
pallescens, tamer lani, velox) 
tauricus 

walli [alaschanicus, 
caucasicus, pallescens, 
tamerlani, tauricus, 
velox allocated to 
Pipistrellus savii by 
Kuzyakin, 1950] 
Eptesicus (Rhyneptesicus) 
nasutus group 

nasutus 

Eptesicus (Eptesicus) 
fuscus group 
bottae 
hingstoni 
serolinus (Including 

andersoni, brachydigitus, 
mirza, pachyomus, pollens, 
shirazensis, sinensis) 



Eptesicus 
capensis group 
brunneus 
capensis ( = notius) (Including 

garambae, grandidieri) 
guineensis (Including 

(?) rectitragus) 
melckorum 
somalicus (Including ugandae, 

vansoni, zuluensis) 

tenuipinnis group 
flavescens ( = angolensis) 
rendalli (Including 

faradjius, phasma) 
tenuipinnis (Including ater) 
Pipistrellus 

circumdatus group 
circumdatus 
mordax 
Eptesicus 



serotinus group 

bottae (Including innesi) 
hottentotus ( = megalurus) 

(Including smithi) 
loveni 
platyops 

serotinus (Including 
isabellinus) 



krefftii) 

Pipistrellus (Neoromicia) 
capensis group 
brunneus 

capensis (Including garambae 
grandidieri, notius, matrokd) 
guineensis (Including 

rectitragus) 
melckorum 
somalicus (Including 

ugandae) 

zuluensis (Including vansoni) 
tenuipinnis group 
flavescens (Including angolensis) 
rendalli (Including 

faradjius, phasma) 
tenuipinnis (Including ater) 
Pipistrellus (Arielulus) 
circumdatus 
cuprosus 
societatis 
Eptesicus 

Eptesicus (Eptesicus) 
nilssonii group 
bobrinskoi 
gobiensis (Including 

centrasiaticus, kashgaricus) 
nilssonii (Including japonensis, 

(?) parvus; propinquus) 
nasutus group 

nasutus (Including batinensis 
matschiei, pellucens, 
walli) 



serotinus group 
serotinus subgroup 

bottae (Including anatolicus, 

hingstoni, innesi, ognevi, 

omanensis) 
brasiliensis (Incluidng andinus, 

argentinus, chiriquinus, 

melanopterus) 
diminutus (Including 

dorianus , fide Us) 
furinalis (Including inca, 

montosus) 
fuscus (Including hispaniolae, 

peninsulae) 
guadeloupensis 
hottentotus ( = megalurus) 



VESPERTILIONINE SYSTEMATICS 



283 



Table 3-cont. 



Tate( 19420) 



Koopman(1973, 1975) 



Hill & Harrison 



sodalis (Including 
ognevi) 



demissus group 

demissus 

Eptesicus (Pareptesicus) 
pachyotis group 
pachyotis 



flower i group 
floweri ( = lowei) 



(Including smithi) 
innoxius (Including 

punicus) 
loveni 
lynni 

serotinus (Including andersoni, 
brachydigitus , horikawai, 
inter medius, isabellinus, 
mirza, pachyomus, pollens, 
pashtomus, platyops, 
shirazensis, sinensis, 
sodalis, turcomanus) 
tatei 
demissus subgroup 

demissus 
(?) pachyotis subgroup 

pachyotis 

Eptesicus (Rhinopterus) 
floweri (Including lowei) 



284 



J. E. HILL & D. L. HARRISON 



base 





shaft 





1 I 








Fig. 1 Bacular types in Pipistrellus and Eptesicus (see text). Scale a-c = 0-5 mm; d-g = 1 mm. 



VESPERTILIONINE SYSTEMATICS 



285 









Fig. 2 Baculum of a, Pipistrellus pipistrellus (D, LL, reversed); b, P. nathusii (D, RL, RVL); c, P. 
papuanus (D, RL); d, P. subflavus (D, RL); e, P. drcumdatus (D, LL, RVL). Scale = 0-5 mm. 



286 



J. E. HILL & D. L. HARRISON 









Fig. 3 Baculum (D, RL) of a, Pipistrellus abramus; b, P. endoi; c, P. paterculus; d, P. ceylonicus (raptor). 

Scale = 2 mm. 



VESPERTILIONINE SYSTEMATICS 



287 










mm* ......,,... .' 

^V.MtV.V.Jm...->-^' 




Fig. 4 Baculum (D, RL) of a, Pipistrellus babu; b, P. collinus; c, P. murrayi; d, P. angulatus (ponceleti). 

Scale = 1 mm. 




^.......:n.<^ v ,,.:... ; :... 1 ......,.,. " w " r '^ > 




Fig. 5 Baculum (D, RL) of a, Pipistrellus kuhlii; b, P. maderensis; c, P. deserti; d, P. rusticus. 

Scale = 0-5 mm. 



VESPERTILIONINE SYSTEMATICS 



289 










Fig. 6 Baculum of a, Pipistrellus savii (D, RL); b, P. HO/IMS (D, RL); c, P. rusticus (D, RL); d, P. helios 
(D, RL); e, P. anchietae (D, LVL, reversed). Scale = 0-5 mm. 








S?>t^KW^^ .,,,...-,.,.:;:.,. , : -..,.. wmc.r.l.UW 









Fig. 7 Baculum of a, Pipistrellus arabicus (D, RL, RVL); b, P. coromandra (tramatus) (D, RL); c, P. 
coromandra (D, RL); d, P. ceylonicus (D, RL); e, P. crassulus (D); f, P. nanulus (D, RL); g, P. mimus 
(D, RL); h, P. stenopterus (D, RL, RVL). Scale = 1 mm. 













Fig. 8 Baculum of a, Pipistrellus affinis (D, RL); b, P. peter si (D, RL); c, P. pulveratus (D, RL, RVL); d, 
P. Hesperus (D, LL, reversed, LVL); e, P. kitcheneri (D, RL, RVL); f, P. lophurus (D, RL); g, P. 
tasmaniensis (D, RL, V). Scale = 1 mm. 




Fig. 9 Baculum (D, RL except where stated) of a, Pipistrellus imbricatus; b, P. macrotis; c, P. societatis, 
d, P. tennis (nitidus) (D, RL, RVL); e, P. anchietae ('Vesperus' bicolor); f, P. bodenheimeri; g, P. 
eisentrauti; h, P. cuprosus. Scales a-g= 1 mm; h = 0-5 mm. 









_l 








Fig. 10 Baculum (D, RL) of a, Pipistrellus rueppellii (pulcher); b, P. rueppellii; c, P. adamsi; d, P. 
westralis; e, P. javanicus; f, Nyctalus noctula; g, P. wattsi; h, P. mackenziei (c, d, g, h from Kitchener 
etal, 1986). Scales = a, b, e, f=2mm;c, d, g, h= 1 mm. 



294 



J. E. HILL & D. L. HARRISON 














Fig. 1 1 Baculum (V, RL) of a, Pipistrellus pumilus pumilus; b, P. pumilus (caurinus); c, P. vulturnus; d, P. 
douglasorum; e, P. regulus; f, P. sagittula (a-c, e, f from McKean et al., 1970; d from Kitchener, 1976). 
Scale = 2 mm. 













Fig. 12 Baculum (D, RL) of a, Pipistrellus capensis (matrokd); b, P. capensis; c, P. guineensis; d, P. 
zuluensis; e, P. rendalli (with anterior view); f, P. melckorum; g, P. capensis; h, P. somalicus', i, P. 
capensis ('minutus'); j, P. tenuipinnis; k, P.pumilus. Scale = 1 mm. 












Fig. 13 Baculum (D, RL) of a, Eptesicus fuscus; b, E. hottentotus (megalurus); c, E. furinalis; d, E. 
brasiliensis (andinus); e, E. bobrinskoi; f, E. floweri; g, E. serotinus; h, E. serotinus (isabellinus); i, E. 
fuscus (hispaniolae); j, . bottae (innesi); k, . brasiliensis: 1, E. flower i(lowei). Scale = 1 mm. 



VESPERTILIONINE SYSTEMATICS 



297 









Fig. 14 Baculum (D, RL except where stated) of a, Eptesicus bottae (omanensis); b, Pipistrellus rendalli 
(? brunneus); c, Eptesicus nasutus; d, Plecotus teneriffae (D) (from Ibanez & Fernandez, 1986). 
Scales = 1 mm. 



298 



J. E. HILL & D. L. HARRISON 





Fig. 15 Baculum (D, RL except where stated) of a, Eptesicus nilssonii (D) (from Topal, 1958); b, 
Baeodon alleni (from Brown ef a/., 1971); c, Pipistrellus peguensis (from Sinha, 1969); d, P. camortae; 
e, Idionycteris phyllotis, f, Plecotus townsendii (pallescens), g, P. raftnesquii (e-g from Nader & 
Hoffmeister, 1983; h, P. rafinesquii (macrotis) (from Hamilton, 1949); i, Bauerus dubiaquercus (from 
Pine etal., 1971). Scales a = 0-5 mm; b, e-i = l mm;c, d = 2mm. 




I . I I I I 

Fig. 16 Baculum (D, RL except where stated) of a, Otonycteris hemprichii; b, Philetor brachypterus (D, 
RVL); c, Nyctophilus gouldi, d, Scotozous dormeri; e, Nycticeinops schlieffenii', f, Laephotis wintoni; g, 
Scotorepens balstoni; h, S. greyii; i, Scoteanax rueppellii. Scales a-h = 1 mm; i = 2 mm. 














Fig. 17 Baculum of a, C//a//no/o^w5 wor/o (D); b, C. gouldi (D); c, C. nigrogriseus (rogersi) (D); d, C. 
picatus (D); e, C. tuberculatus (D); f, Lasionycteris noctivagans (D, RL); g, Scotophilus nigrita (gigas) 
(D, RL); h, 5. heathii(D, RL); i, S. kuhlii(D, RL);j, 5. nigritellus (D, RL); k, Nycticeius humeralis (D, 
LL, reversed). Scale = 1 mm. 













Fig. 18 Baculum (D, RL) of a, Glischropus tylopus; b, Antrozous pallidus; c, Histiotus velatus; d, H. (?) 
macrotis; e, ^. macrotis; f, Dasypterus argentinus; g, Scotomanes ornatus; h, Tylonycteris pachypus; i, 
r. robustula;j, Barbastella barbastellus; k, Rhogeessa tumida. Scale = 1 mm. 













I 

Fig. 19 Baculum of a, Glauconycteris poensis (D); b, G. variegata (D); c, G. beatrix (D); d, G. argentata 
(D); e, G. humeralis (D); f, G. variegata (papilio) (D); g, Plecotus auritus (D); h, /*. austriacus (D); i, 
Myotis ridleyi (D, RL); j, M. nattereri (D, RL); k, Pizonyx vivesi (D, RVL); 1, Lasiurus cinereus (D, 
RVL). Scale = 1mm. 











Fig. 20 Baculum (D, RL) of a, Scotoecus albigula; b, 5. hindei (falabae) c, 5. hirundo; d, S. hindei; e, S. 

albofuscus. Scales a-d = 2 mm; e = 1 mm. 









d %**^ 








I I 




Fig. 21 Baculum (D, RL except where stated) of a, Hesperoptenus tomesi, b, H. tickelli, c, H. doriae (a-c 
from Hill, 1 976); d, la io (D) (from Topal, 1 970); e, Scotorepens orion, f, S. sanborni (e, f from Kitchener 
& Caputi, 1985); g, Hesperoptenus blanfordi (from Hill & Francis, 1984); h, Scotoecus pallidus (from 
Agrawal & Sinha, 1973); i, Vespertilio murinus (V, RL) (from Topal, 1958); j, V. orientalis (from 
Wallin, 1969). Scales a-c = 2 mm; d-h, j= 1 mm; i = 0-5 mm. 







_- ' - 











Fig. 22 Baculum (D, RL) of a, Nyctophilus bifax; b, AT. geoffroyi (pallescens); c, M microtis; d, Af. gouldi; 
e, A 1 , geoffroyi (pacificus); f, A 7 ^. gouldi (sherrini); g, A 7 , daedalus; h, Pharotis imogene. Scale = 2 mm. 






British Museum (Natural History) 

The birds of Mount Nimba, Liberia 

Peter R. Colston & Kai Curry-Lindahl 



For evolution and speciation of animals Mount Nimba in Liberia, Guinea and the Ivory Coast is 
a key area in Africa representing for biologists what the Abu Simbel site in Egypt signified for 
archaeologists. No less than about 200 species of animals are endemic to Mount Nimba. Yet, this 
mountain massif, entirely located within the rain-forest biome, is rapidly being destroyed by 
human exploitation. 

This book is the first major work on the birds of Mount Nimba and surrounding lowland 
rain-forests. During 20 years (1962-1982) of research at the Nimba Research Laboratory in 
Grassfield (Liberia), located at the foot of Mount Nimba, scientists from three continents have 
studied the birds. In this way Mount Nimba has become the ornithologically most thoroughly 
explored lowland rain-forest area of Africa. 

The book offers a comprehensive synthesis of information on the avifauna of Mount Nimba 
and its ecological setting. During the 20 years period of biological investigations at Nimba this in 
1 962 intact area was gradually opened up by man with far-reaching environmental consequences 
for the rain-forest habitats and profound effects on the birds. Therefore, the book provides not 
only a source of reference material on the systematics, physiology, ecology and biology of the 
birds of Mount Nimba and the African rain-forest, but also data on biogeography in the African 
context as well as conservation problems. Also behaviour and migration are discussed. At 
Nimba a number of migrants from Europe and/or Asia meet Afrotropical migratory and 
sedentary birds. 

Professor Kai Curry-Lindahl has served as Chairman of the Nimba Research Laboratory and 
Committee since its inception in 1962. Peter Colston is from the Subdepartment of Ornithology, 
British Museum (Natural History), Trmg, and Malcolm Coe is from the Animal Ecology 
Research Group, Department of Zoology, Oxford. 

1986, 129pp. Hardback. 565 00982 6 17.50. 






Titles to be published in Volume 52 



Miscellanea 

A revision of the Suctoria (Ciliophora, Kinetofragminophora) 5. The Paracineta 
and Corynophora problem. By Colin R. Curds 

Notes on spiders of the family Salticidae 1. The genera Spartaeus, Mintonia and 
Taraxella. By F. R. Wanless 

Mites of the genus Holoparasitus Oudemans, 1936 (Mesostigmata: Parasitidae) in 
the British Isles. By K. H. Hyatt 

The phylogenetic position of the Yugoslavian cyprinid fish genus Aulopyge Heckel, 
1841, with an appraisal of the genus Barbus Cuvier & Cloquet, 1816 and the 
subfamily Cyprininae. By Gordon J. Howes 

Revision of the genera Acineria, Trimyema, and Trochiliopsis (Protozoa, 
Ciliophora). By H. Augustin, W. Foissner & H. Adam 

The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) with a 
systematic review, a synopsis of Pipistrellus and Eptesicus, and the descriptions of 
a new genus and subgenus. By J. E. Hill & D. L. Harrison 

Notes on some species of the genus Amathia (Bryozoa, Ctenostomata). 

By P. J. Chimonides 



Printed in Great Britain by Henry Ling Ltd.. at the Dorset Press, Dorchester, Dorset 



Bulletin of the 

British Museum (Natural History) 



Notes on some species of the genus Amathia 
(Bryozoa, Ctenostomata) 



P. J. Chimonides 



Zoology series Vol 52 No 8 27 August 1987 



The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four 
scientific series. Botany Entomology, Geology (incorporating Mineralogy) and Zoology, and 
an Historical series. 

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World List abbreviation: Bull. Br. Mus. nat. Hist. (Zool.) 



Trustees of the British Museum (Natural History), 1987 



The Zoology Series is edited in the Museum's Department of Zoology 

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ISBN 0565 05032 X 

ISSN 0007 1 498 Zoology series 

Vol 52 No. 8 pp 307-358 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD Issued 27 August 1987 



Notes on some species of the genus A mat hi; 
Ctenostomata) 

P. J. Chimonides 

Department of Zoology, British Museum (Natural History), Cromwel 



Contents 

Synopsis ......... 

Introduction 

Definition and assessment of taxonomic characters 

Materials and methods 

Key to species 

Systematic section 

Discussion 

References 




Synopsis 

The species of the genus Amathia have presented difficulties of recognition for a long time. Even the most 
recent revisionary accounts have failed to establish the separate identity of many of the species; these have, 
instead, been submerged in erroneous synonymies. Fifteen species are dealt with in full. In the main, species 
without significant spiralling of the autozooid groups are considered. Three new species are introduced: A. 
guernseii, A. intermedis and A. tricornis. A fourth species, A. populea Busk MS in d'Hondt, is recognised as 
new. A. distans var aegyptana is raised to specific rank. A. cornuta Lamouroux (preocc.) is renamed A. 
lamourouxi. A. obliqua and A.plumosa MacGillivray are redescribed. A. alternata Lamx., A. biseriata Krauss, 
A. brongniartii Kirkpatrick, A. lendigera Linn., A. pruvoti Calvet, A. pinnata and A. wilsoni Kirkpatrick, A. 
woodsii Goldstein, are redefined with type material selected. Characters for use in taxonomic and systematic 
discrimination are introduced, and brief comment is made on the zoo- and palaeo-geography of the species 
dealt with. 



Introduction 

The genus Amathia was erected in 1812 by J. V. F. Lamouroux, consequential to the study of 
material collected from Tasmania and the south coast of Australia by C. A. Lesueur and F. Peron, 
during the years 1800-1804 (Tenison Woods 1880, d'Hondt 1979). 

However, Sertularia lendigera of Ellis (1755) from European waters became the type species of 
the genus, by virtue of being Lamouroux's sole mentioned species at the introduction of the genus 
(I.C.Z.N. article 69(d)). The species itself was validated with the publication of the 10th edition of 
the Sy sterna Naturae by Linneaus in 1758 (and is therefore technically that of Linneaus). 

Ryland (1982) gave a revised perspective classification of the genus but there are differences 
between his definitions of higher categories, including Amathia, and the characters of the genus 
presented here. At the Family level, Ryland described the 'zooids' as being 'radially symmetrical, 
no face being partly membranous'; and at the Superfamily level, he described 'branching being 
irregular'. Both descriptions are inaccurate for Amathia. Similarly d'Hondt (1983) for the Family 
level, also described 'External autozoecial symmetry' as 'radiated', while at the Superfamily level, 
there was some ambiguity in the definitions of the characteristics employed e.g. for the 'Zoarium 
. . . autozoecia unconnected to their neighbours'. Clearly there is need for a review of the characters 



Bull. Br. Mus. nat. Hist. (Zool.) 52 (8): 307-358 



Issued 27 August 1987 



307 



308 P. J. CHIMONIDES 

used in the definition of the higher taxonomic levels, although this is beyond the scope of this 
account. 

The persistent problem has been how to differentiate between the numerous species. The brief 
descriptions often given are of little help, and in fact have led to some confusion. Often, widespread 
geographical distributions have been suggested. Where no figures or specimens are available, it is 
unlikely that the true identity of some species will ever be recognised. Despite the efforts of d'Hondt 
(1979, 1983) the picture still remains clouded. 

This account attempts to establish criteria for species differentiation within the genus; to identify 
some species groupings based upon these criteria and in the process to discuss and correct past 
misconceptions. 

In general, the species of Amathia may be assigned to either of two groupings: those with 
autozooids spirally disposed about the stolon; those with autozooids disposed linearly along the 
stolon. It is mainly the latter group which is discussed here. Where spirally disposed species are 
dealt with, this is mainly to obviate possible confusion with those species (i.e. A. alternata and 
A. pruvoti) in which marked twisting of the autozooid groups occurs along the stolon. It is in the 
context of comparison with A. pruvoti that A. distans var aegyptana is considered. 

Definition and Assessment of Taxonomic Characters 

Waters (1910) in his brief account of the genus, outlined a number of characters which may serve as 
a foundation on which to build an understanding of both the genus and its species. These charac- 
teristics may be added to, and arranged in what is considered here to be an order of decreasing 
reliability, reflecting an increase in their intraspecific variability. 

List of species discrimination characteristics in order of reliability 

( 1 ) budding pattern of stolons 

(2) development of any kenozooidal processes or rhizoids and their orientations 

(3) arrangement of autozooids about the stolons 

(4) autozooidal thickening 

(5) profile of autozooids and stolons 

(6) number of autozooids and proportion of stolon occupied by autozooids 

(7) dimensions of components 

The potentially informative characters of larval type, larval metamorphosis, ancestrula 
formation and initial colony development are generally not known and hence cannot be evaluated. 
Despite the explicit account given by Barrois (1877) for A. lendigera, the ancestrula and earliest 
astogenetic stages, for example, have not been recognised in any of the specimens examined. In 
some specimens it was clear that this part of the colony was absent; in others, it was impossible to 
see because of heavy overgrowth by the colony's own rhizoids or by spatial competitors. For these 
same reasons, in the following systematic accounts, no information is given on the non-erect part 
of the colony for the majority of species. It is possible that some colonies are the result of associ- 
ation between the products of more than one ancestrula, (without the necessity for fusion to 
have occurred, especially in the non-arborescent growth forms). Zimmer and Woollacott (1977a) 
suggested that the larval type of all stoloniferan ctenostomes is the same. It would appear however, 
that their conclusions were drawn from only three species: 'Amathia lendigera', Bowerbankia 
pustulosa (Ellis and Solander) and Zoobotryon verticillatwn (Delle Chiaje). Furthermore, Zimmer 
and Woollacott (19776) pointed out that past accounts of metamorphosis of larvae in this group 
were inconsistent, and that 'additional work is essential to clarify the pattern(s) of metamorphosis' 
of the larval type. Waters' (1910) intuitive suggestion of the 'valuable assistance' which the primary 
zooecia might give must, therefore, be discounted for the present. 

Extensive examination of several large colonies indicates that the branching pattern of stolons 
remains remarkably consistent within species. Differences in branching pattern may be inferred to 
have been microenvironmentally induced in that they tend to be sporadic, involve the development 
of new stolens from astogenetically early regions of the colony, and are often associated with the 



AMATHIA 309 

presence of epibionts. The budding patterns, together with kenozooidal processes, rhizoids, auto- 
zooidal thickenings and disposal of autozooids about the stolons, can give rise to characteristic 
colony shapes which, with familiarisation, allow identification of species by casual inspection. 

The growth of a colony relies essentially on the production of the supporting 'stolonal' keno- 
zooids. Where such a kenozooid is destined to bear autozooids, the autozooids are usually seen to 
develop at about the same time as the kenozooid lengthens through apical growth. Autozooid- 
bearing kenozooids are here termed stolons. The position of the autozooids and the proportion of 
stolon occupied by them is highly regular. Stolonal and autozooidal growth is considered to have 
ceased with the production of septa at the distal end of the stolon, and the subsequent appearance 
of daughter stolonal buds. The kenozooidal processes of character No. 2 in the above list appear to 
be growth-terminating features. These are usually distinguishable from potential autozooid- 
bearing stolonal kenozooids by being straighter, narrower, often tapering to a point, and 
frequently being subdivided by septa. 

Where rhizoids are to be produced by a structure, the origin of each rhizoid is marked first by the 
appearance of an oval window in the cuticle. Rhizoids then develop as papilliform outgrowths of 
these windows, proximally directed along the colony, growing towards the colony base. The 
rhizoids sometimes fuse with each other en route, overgrowing and obscuring underlying stolons, 
and forming a trunk-like mass. Autozooids and lateral branches of overgrown stolons are often 
shed. When rhizoids are produced, the resulting colony form is usually arborescent. 

The overall cuticular thickening of any colony appears uniform, except at the regions of the 
growing tips, where it is thinner. This level of cuticular thickening can differ between colonies of the 
same species. As this thickening is contributory to colour, it follows that colour is also variable. 
Within all species, there are localised areas of thickening, which tend to be constant. Thickening in 
the autozooids, which gives their groupings a characteristic appearance, may be used to discrimi- 
nate between species. Two conditions occur: one where the walls between autozooids are differen- 
tially thickened (inner-wall thickening); the other where the outer walls are differentially thickened 
(outer-wall thickening) (see Fig. IB, C). 

The arrangement of autozooids on the stolon is usually described as being paired, or as a biserial 
row. Although this appears correct, in all specimens examined, displacement of autozooids occurs, 
so that the autozooids of one row interlock with the recesses between the autozooids in the other 
row (Dalyell, 1847 for A. lendigerd). Very frequently, this emphasises a single proximal-most 
autozooid in each group. No pairing of autozooids may be confidently assigned throughout a 
colony in any species, and there can be odd or even numbers of autozooids in any autozooid group. 
In some cases, notably those with inner-wall thickening, the proximal-most autozooid tends to be 
larger in cross section and displaced centrally, such that it can be very difficult to assign it to a row 
of origin at any stage in its ontogeny. 



Materials and methods 

Specimens used for study were mainly those of the British Museum (Natural History), London, 
(BMNH) and The Manchester Museum, (MM), with additional material referred to as follows: 
Laboratoire de Biologic des Invertebres Marins et Malacologie of the Museum National 
d'Histoire naturelle, Paris, (LBIMM); the National Museum of Victoria, Melbourne, (NMV); 
the Rijksmuseum van Natuurlijke Historic, Leiden (RM); and the U.S. National Museum, 
Washington, (USNM). 

Often, the material for study had been preserved dried, with resulting distortions. To observe 
the autozooidal characteristics preserved in the cuticular thickenings, it was found far better to 
rehydrate the specimens although it was still possible to make identifications without treatment. 
Rehydration was carried out using tri-sodium phosphate in 7-10% aqueous solution, with subse- 
quent transfer to distilled water and then via a sucession of increasing concentrations of alcohol, to 
80% concentration for storage. From this process, specimens regained the turgidity associated 
with their living state. It was from specimens in this turgid state that measurements were taken. 



310 P. J. CHIMONIDES 

Some care was needed, as rupture of specimens was possible through the initial high osmotic 
differential established on transfer to distilled water. In some cases, specimens failed to reflate 
because of existing ruptures in their cuticles. 

In the ensuing descriptions, anterior is used to denote the side at that location bearing auto- 
zooids, and posterior, the side opposite (see Fig. ID, E). Dimensions are given in millimetres and 
are means of a minimum of 30 measurements. Where shape negated the validity of a single sample 
measurement, extreme dimensions are given, these also being the means of 30 measurements each. 
Measurements were made in ontogenetically complete components, near distal regions, avoiding 
where possible, astogenetically earlier (older) regions of the colony. No attempt was made to 
determine intra-colony variations quantitatively. Where these were noted, they were assessed 
subjectively. 
The following abbreviations are used: 

SI. length of stolon 

Sd. diameter of stolon, at location specified; usually midway along the proximal autozooid-free end. 

Zh. autozooid height to the highest point on the rim of thickening, of tallest autozooids, unless otherwise 

specified. 

Zw. autozooid width, measured along the stolonal axis. 

Z/S. the linear proportion of stolon occupied by autozooids 

Zn. the number of autozooids per autozooid group (and apparent number of 'pairs') 

Tpl. length of terminal process 



Key to species 

(Identification is best attempted with plentiful material.) 

1 Rhizoids developed, colony frequently aborescent 

Rhizoids not developed, colony not arboresecent, no terminal processes and Z/S ratio < 50% . 1 3 

2 Autozooidal thickening inner-wall brongniarttt 

Autozooidal thickening outer-wall 

3 Branching nearly always bifurcate 4 

Branching primarily tri- and tetrafurcate, bifurcation may also be present 10 

4 Terminal processes developed 5 

Terminal processes not developed 

5 A pair of lanceolate, single-kenozooidal terminal processes developed at the distal end of each 

autozooid group, arising in the same direction as the autozooids . . . lamourouxi 
Lanceolate terminal processes of compound kenozooidal construction, each filament developed in 
place of a normal stolon, sometimes branched 6 

6 Rhizoids developed postero-laterally, terminal processes never branched . . . populea 
Rhizoids developed anteriorly, terminal processes often forked woodsii 

1 Autozooids re-orientated by approx. 180 deg. from stolon to stolon, polyrhizoidy (see page 335) 

possible alternata 

Autozooid orientation from stolon to stolon maintained within 30 deg., rhizoids paired at most . 8 

8 Rhizoids developed anteriorly, autozooids with marked distal inclination, autozooid group 

profile diminishing distally guernseii 

Rhizoids developed postero-laterally, autozooid group profile horizontally even i.e. level . . 9 

9 Stolons curved anteriorly, curvature increasing distally, autozooid group arranged in line with 

stolonal axis biseriata 

Stolons straight, autozooid group set obliquely to stolonal axis obliqua 

10 Autozooid-bearing stolons developed laterally from a central axis of stolon-sized, or larger, 

kenozooids. Autozooid-bearing stolon sequences end with compound terminal processes, these 

often forked plumosa 

Autozooid-bearing stolons developed laterally from a central axis of other autozooid bearing 

stolons 11 



AMATHIA 311 

1 1 Central axis stolons undergo trifurcation only; indistinguishable from lateral stolons . pinnata 
Central axis stolons usually undergo tetrafurcation, a fourth autozooid-bearing stolon 

developed posteriorly. Central axis stolons morphologically distinguishable from lateral 
stolons, differences may be slight 12 

12 Autozooid-bearing stolon sequences end with lanceolate, compound kenozooidal, terminal pro- 

cesses, each replacing a normal stolon and thus in complements of three. Pronounced difference 

between central axis and lateral stolons tricornls 

Autozooid-bearing stolon sequences with pinnate, compound terminal processes; each assemb- 
lage replacing stolons in other, regular positions, giving characteristic arched colony sub units. 
Difference between central axis and lateral stolons slight wilsoni 

13 Autozooid groups regularly twisted along stolon length pruvoti 

Autozooid groups rarely showing any twist 14 

14 Stolons often in rectilinear series, straight, sometimes undergoing trifurcation. Autozooid groups 

often remote from subsequent branching point. Autozooids usually erect . . intermedis 

Stolons of variable length, usually short, sculptured and posteriorly deflected. Autozooid groups 

overlie subsequent branching point, autozooids inclined distally, the lean increasing distally . 

lendigera 



312 P. J. CHIMONIDES 

Systematic Section 

Phylum BRYOZOA Ehrenberg, 1831 
Class GYMNOLAEMATA Allman, 1856 

Order CTENOSTOMATA Busk, 1852 
Genus AM A THIA Lamouroux, 1812: p. 184 

Part Sertularia Linnaeus, 1758. 

Serialaria Lamarck, 1816. 

Part Valkeria Dalyell, 1847. 

AmathellaGray, 1858. 

CharadellaGray, 1858. 

Serialia Gray, 1858. (errorum pro Serialaria Lamarck, 1816). 

SpiraliaGray, 1858. 

CornaliaGray, 1858. 

Amathia: Bobin & Prenant, 1956: (incomplete cum. syn., NB. Gray 1858 misquoted as 1848); Ryland, 1982; 

Winston, 1982;d'Hondt, 1979, 1983;Hayward, 1985. 

TYPE SPECIES. A. lendigera (Linnaeus 1758 sensu Ellis 1755) Lamouroux 1812: p. 184. 
GENERIC DESCRIPTION. Colonies mainly erect with a creeping base, this sometimes extensive. 
Autozooid groups displaced towards the distal portion of the stolon. Stolons may produce 
rhizoids, proximally disposed. Distal, mainly growth-terminating kenozooidal processes may be 
developed from various positions. Autozooids with gizzards, borne on kenozooidal stolons, 
arising from rosette plates, in groups, connate for at least part of their length, appearing biserially 
arranged as a straight or spiral series. 

REMARKS. The only attempt to regroup species comprising the genus Amathia was made by 
Gray (1858, duplicated 1859). Gray introduced several indeterminate subgeneric or generic 
groups, the type species of which were insufficiently described and not illustrated. The great 
majority of the limited characteristics employed are variable within species, such that none of the 
divisions Gray introduced exclusively defines any species group identifiable within the genus. 
Bobin & Prenant (1956) are followed here in assigning all species described to the genus Amathia. 

Amathia lendigera (Linnaeus, 1758) 
(Figs 6A, 7A) 

? Sertularia lendigera Ellis, 1755: 27, pi. 15 (figs 24B, 24b). 

Sertularia lendigera Linnaeus, 1758: 812. 

Amathia lendigera Lamouroux, 1812: 184. 

Not Amathia lendigera: MacGillivray, 1895: 135, pi. B (fig. 1). 

Not Amathia lendigera: O'Donoghue & de Watteville, 1944: 430 (= A. populea). 

Part Amathia lendigera: Bobin & Prenant, 1956: 280. 

Amathia lendigera: Hay ward, 1985: 134, fig. 45B. 

MATERIAL EXAMINED 
Neotype (selected here): BMNH; 1963.1.8.3, Chichester Harbour, H. G. Stubbings collected. 

OTHER MATERIAL 

BMNH; 1827.11.18.8, no locality. 1852.3.16.62, Weymouth. 1882.7.7.85, no locality. 1887.7.23.5, Solent, 
I.O.Wight. 1891.8.7.18, Portland, Dorset. 1897.8.9.67, Weymouth Bay, Portland, 10 fthms. (18.29m). 
1899.5.1.21 l,?OffSaints Bay, Guernsey? 1900.10.30.10-11, Weymouth. 1912.12.21.681, Plymouth. 

MM; 7093-4, Naples. 7095, Roscoff, France. 7096-9, Swanage. 7105, Naples. 7106, St. Raphael, S. France. 
7107, Rapallo( = Rapolla, Italy?). 

DESCRIPTION. Colonies tend to have to have a moderately extensive creeping component of 
stolonal kenozooids. These are adpressed to the substratum and closely follow its profile, showing 
reduced branching in some places and multiple branching in others. These stolonal kenozooids are 
usually of irregular form and length, and only rarely bear autozooids. Bilateral palmate processes 
are often produced, through which adhesion to the substratum is effected. Erect components may 



AMATHIA 313 

be produced at any branching point, with or without continuation of the creeping component. The 
erect components develop as the characteristic autozooid-bearing stolons, arranged in the typical 
form of an orbicular mass, cotton-wool like in appearance, utilizing well the available free space 
near to the substratum. These erect components appear tangled, but are rarely so. Any erect 
component may resume the creeping habit on contact with the substratum. Branching in the erect 
part of the colony is practically always bifurcate, ranging from equally dichotomous to almost 
rectilinear with side branches, these appearing on alternate sides. Bifurcations typically form an 
angle of 90 deg. Autozooid group orientation about the stolon is not usually preserved from stolon 
to stolon. Daughter stolons often arise deflected anteriorly to maternal stolons. Autozooid groups, 
with relatively few autozooids, occurring at the extreme distal end of stolons, frequently over- 
lapping the subsequent branching point. Stolons are often deflected posteriorly at the proximal end 
of the autozooid group, and also raised slightly on the anterior surface at this same region. Stolons 
may be of variable length. Autozooids are outer-wall thickened. Autozooid profile diminishes 
distally, due in part to decreasing autozooid height, and in part to increasing distal inclination of 
the autozooids. A proximal-most autozooid is usually prominent in each group and is displaced 
centrally. Where not truly central, this autozooid remains on the same side of its stolon as the 
direction in which that stolon was budded. The arrangements of autozooids on sister stolons are 
therefore mirror images of each other (see Fig. 5B). Where stolons form linear sequences, auto- 
zooid groups borne on such series tend to show an alternate sequence of autozooid displacements 
on successive stolons. Sometimes sister stolons carry identical autozooid displacements, these 
being opposite to that on their maternal stolon. No overall pattern is discernible within the colony 
in the occurrence of this second state of succession (see Fig. 5C). Rhizoids are absent. 

SI. 1.25-2.75 Z/S. 25-50% 

Sd. 0.75-0.97 Zn. 8-1 7 (appearing as 4^8 'pairs') 

Zh. 0.33-0.50 

Zw. 0.10-0.12 



REMARKS. According to Harmer (1931), the original specimens described and figured by Ellis 
(1755) were not kept (I.C.Z.N. article 73(b) (i), recommendation 69B). Harmer stated that speci- 
mens of A. lendigera were sent to Linnaeus by Ellis, but some 12 years after the publication of the 
nomenclaturally significant 10th edition of Linnaeus' Sy sterna Naturae (1758). Two specimens, 
under the original name ofSertularia lendigera, are still in the collections of the Linnean Society of 
London (Nos. 1298.17 and 1298.18). The specimens are preserved pressed dry on paper, and both 
are labelled as 'lendigera' in Linnaeus' handwriting. From examination of these specimens, some 
doubt arises that Ellis and Linnaeus were sufficiently rigid in their interpretation of A. lendigera. 
Two species are present: specimen 1298.17 is identifiable as A. semiconvoluta (see pages 335, 338); 
while specimen 1298.18 is probably A. lendigera. Linnaeus (1758) has trustingly used Ellis' (1755) 
description verbatim. If the specimens originated from Ellis, Linnaeus may also have accepted 
their identity from him. It is possible therefore, that the mistaken identity of 1298.17 could be 
attributed to Ellis; neither man realising the presence of mixed material. 

However, there is some evidence in support of Harmer' s statement that the Linnaean specimens 
are not Ellis' original (1755) material. Linnaeus is reported to have been in the habit of upgrading 
his botanical collections, with the replacement of older specimens by new, 'some of them not 
conspecific by modern taxonomic standards' (Stearn, 1957), a practice which could also have been 
applied to herbarium preparations of 'zoophytes'. In addition, none of the figures of Ellis (1755) 
correspond with either of the Linnean Society specimens, in particular specimen 1298. 18. Features 
of importance are: the arborescent and open appearance of the colony shape in figure '24b'; the 
number of autozooids per stolon indicated by the magnified view in figure '24B'. Although only a 
single line of autozooids is drawn in the latter figure, this may be interpreted as showing either: a 
single proximal-most autozooid with indications of the outlines of subsequent 'paired' autozooids; 
or possibly a line of 'all paired' autozooids. The condition depicted is readily seen in many dry 
preserved specimens, where only the thickened outer walls, forming the periphery, survive well. As 



314 P. J. CHIMONIDES 

such, 1 7 or 1 8 autozooids would be represented on three of the five stolons; 1 5 autozooids would be 
represented on one of the remaining two; and there is an inexplicable absence of autozooids on the 
remaining fifth and final stolon. Although notionally possible, it is very unusual for A. lendigera to 
show as many autozooids per stolon in direct succession in a colony. The importance of this 
analysis is that figure '24B' is claimed as an exact microscope drawing. 

Harmer (1931) suggested that figures of Ellis be regarded as the lectotype of the species. 
However, the figures are inadequate, no rhizoids are shown, and their presence or absence is not 
indicated in the description. On the cumulative evidence (see above), figures '24b' and '24B' could 
thus be depictions of A. intermedis or even A. guernseii. 

Selection of a neotype specimen is the only satisfactory way to resolve the identity of A. 
lendigera; particularly important as the species is the type of the genus. There is no indication that 
the Linnean Society specimen 1298.18 formed any basis for the description for the species. In 
addition to the uncertainties surrounding its status, 1298.18 unfortunately also lacks sufficient 
locality data, is not in an adequate state of preservation, and so should not be considered. Specimen 
BMNH 1963.1.8.3 is therefore selected here as neotype. It is preserved in alcohol, growing on 
Halidrys siliquosa as is the Linnean Society specimen. BMNH 1963.1.8.3 is erroneously listed by 
d'Hondt (1983) as A. pruvoti, a very different species (see pages 336, 337). 

There is great similarity between A. lendigera and A. intermedis and both resemble A. guernseii 
(see pages 316, 317). The morphologies of all three may overlap in different parts of the colony. A. 
lendigera differs from A. intermedis in that: it tends to have fewer autozooids per autozooid group; 
the autozooids have an increased distal inclination; the autozooid group profile diminishes distally 
more rapidly; the autozooid groups and subsequent bifurcation sites are more condensed relative 
to each other; it has a more compact colony form, with low incidence of rectilinear succession. 
Great care is needed to distinguish between the trifurcation that may occur in the erect part of the 
colony of A. intermedis, and the multiple branching, including trifurcation, which occurs in the 
immediate vicinity of the non-erect part of A. lendigera, as detachment from the substratum is 
frequent in preserved specimens. Non-erect stolons may usually be identified by the nearby 
presence of palmate processes (see Fig. 8A), and the irregular morphology associated with the 
creeping mode. 

The displacement of the proximal-most autozooids in maternal and daughter stolons, may 
reflect the timing of the production of daughter stolons relative to each other. The mirror image 
arrangement (see Fig. 5B) possibly results from the simultaneous production of the daughters. 

Most of the published records for A. lendigera are listed in a lengthy synonymy by Bobin and 
Prenant (1956). However, many of these records are unsupported by specimens available for 
examination and are thus equivocal. In addition, the account these authors give mentions the 
occurrence of rhizoids, and thus includes another species, probably A. guernseii. 

Three specimens in the Waters Collection in the MM. (7100, 7101, 7102) from Zanzibar, are 
superficially similar to A. lendigera. However, notwithstanding the little material present, it is 
possible to see that the autozooid groups lack any characteristic distal inclination. In addition, the 
linearly disposed stolons seem to be arranged in true rectilinear fashion and lack any posterior 
deflection associated with stolons of their length as in A. lendigera. Another specimen (7104) from 
Menton (southern France) labelled 'A. lendigera', shows trifurcation at four stolons in almost 
direct succession, but conforms in most other characteristics. These stolons are all in proximity to 
substratum attachment sites and are probably not typical of the whole colony budding pattern. 
These is not enough material to be certain about this or the true identity of the specimen. The 
locality is, however, within the expected distribution area of A. lendigera. Some of MacGillivray's 
specimens (NMV 65387-8) marked 'British', are A. lendigera. Additional material (NMV 
65383-5) labelled 'A. lendigera' and from Western Port, Australia, is a different species. These 
specimens bear little resemblance to the 'British' material, and in addition, show evidence of 
rhizoids. Where the rhizoids are not obvious, careful illumination is required to observe the oval 
window precursors. The specimens are probably early astogenetic stages of A. lamourouxi, but 
there is not enough material to be certain; the characteristic terminal processes are not present, and 
the identity is inferred from the branching characteristics. The 'Australian' specimens may be the 



AMATHIA 315 

material described as A. lendigera by MacGillivray (1895, pi. B, fig. 1), although the actual 
specimen figured does not appear to have been recorded. 

DISTRIBUTION. A. lendigera is known from the Thames estuary, and along the south and west coasts 
of England. The species also occurs off the north coast of Africa, off Mediterranean southern 
France, and Naples and 'Rapallo' in Italy. Substrata recorded are rocks and the alga Halidrys 
siliquosa. 

Amathia intermedia sp. nov. 
(Figs. 6C, 7C) 

? Serialaria lendigera: Johnston, 1838: fig. 40. 

? Serialaria lendigera: Johnston, 1847: fig. 68. 

? Serialaria lendigera: Couch, 1844: pi. 16. 

Valkeria lendigera Dalyell, 1847: 249, pi. 52 (fig. 2). 

? Part Amathia lendigera: Bobin & Prenant, 1956: fig. 124, 1, IV. 

Holotype: BMNH; 1887.5.2.18 part, Hastings, England. 

Paratypes: BMNH; 1842.12.9.14, Belfast Bay. 1847.9.24.184, North'd ( = Northumberland?) Coast. 
1887.5.2. 18 part, Hastings. 1963.2.10.1, Scarborough. 1985.3.2.1a, Ib, Yarmouth. 1985.3.2.2, Bournemouth. 
1985.3.2.3, no locality. 

ETYMOLOGY. The species at one time seemed intermediate in character between A. lendigera and 
A . guernseii. 

DESCRIPTION. In the erect part of the colony, branching is primarily bifurcate, ranging from equally 
dichotomous to rectilinear series with side branches. There is a disposition to the latter condition, 
where at a bifurcation, one daughter stolon usually remains in line with the main axis of the 
maternal stolon, while the other daughter stolon appears sequentially on alternate sides. These 
lateral daughter stolons are produced at the same distal inclination to the maternal stolon axis as 
the maternal stolon autozooids. Their lateral angular inclination may be from 0-90 deg. to the 
orientation of the maternal autozooids, but usually ranges from 10-30 deg. Occasionally there is a 
trifurcation, in which, of the three daughter stolons produced, the middle one lies in the rectilinear 
position. The other two are produced one on either side, separated from the central one by 
approximately 45 deg. The autozooids on the maternal stolon bisect this angle. Autozooid groups 
occur towards the distal end of stolons, but there is usually a further autozooid-free portion of 
stolon, distal to the autozooid group. This is often axially well divided into small branches, the 
subdivisions orientated in the same direction as, and supporting, the daughter stolons. There is 
frequently a further autozooid-free length between the end of the autozooid group and this region 
of division, approximately equal to the diameter of one autozooid. Stolons are often straight, 
showing little sign of accommodating the autozooids borne. The autozooids tend to be erect, and 
of even height throughout the autozooid group, although autozooid group profile sometimes 
diminishes at the distal end. This is due in part to an increased inclination in the autozooids, and in 
part due to decreasing autozooid height. Autozooids are outer-wall thickened, but thinly so 
overall, and pale yellow brown in colour. Viewed anteriorly, a proximal-most autozooid is usually 
evident in each autozooid group. The occurrence of this autozooid, the pattern of autozooid 
displacements, and the succession states of autozooid displacements on the stolons, are identical to 
that found in A . lendigera (see page 3 1 3). The orientation of the autozooid group about the stolon is 
not always preserved from maternal to daughter stolons; re-orientations of up to 180 deg. may 
occur. No rhizoids are produced, and the colony attains a diffuse cotton-wool like appearance. The 
non-erect part of the colony does not appear as extensive as the erect part. Stolonal kenozooids in 
the non-erect part of the colony: produce branches occasionally; tend not to bear autozooids; are 
not of the same appearance as those of the erect part, in being elongated, sometimes twisted, and 
generally following the profile of the substratum. Erect components may be produced at any 
branching point, these assuming the normal erect growth pattern. Attachment to the substratum is 
effected through lateral palmate processes, often developed bilaterally from the adnate stolonal 
kenozooids. 



316 P. J. CHIMONIDES 

SI. 1.75-3.25 Z/S. 35-50% 

Sd. 0.80 Zn. 8-29 (appearing as 4- 14 'pairs') 

Zh. 0.35-0.45 

Zw. 0.10 

REMARKS. A. intermedis resembles A. lendigera and A. guernseii, the closest similarity being with 
the former. A. intermedis may be distinguished from A . lendigera in having the following character- 
istics: trifurcations in the erect part of the colony; a tendency towards higher numbers of auto- 
zooids in the autozooid groups, and longer stolons; a staggered occurrence of autozooid groups 
and branching sites; a more open colony form, resulting from a higher occurrence of rectilinear 
succession in the stolons. A. intermedis may be distinguished from A. guernseii primarily in the fact 
that A. guernseii develops rhizoids. 

As with A. lendigera, the displacement of the proximal-most autozooids in maternal and 
daughter stolons may reflect the timing of the production of the daughter stolons relative to each 
other (see page 314). 

BMNH 1842.12.9.14, 1847.9.24.184, from Johnston's collection, are A. intermedis, but it is not 
known if any of this is his figured material (1838, fig. 40, 1847, fig 68). 

DISTRIBUTION. The species is known from the east and south-eastern coasts of England, and also 
from Belfast Bay. The only substratum recorded is the alga, Halidrys siliquosa. 

Amathia guernseii sp. nov. 
(Fig 2A, 6B, 7B) 

Holotype: BMNH; 1898.5.7.189, Saints Bay, Guernsey. 

Paratypes: BMNH; 1912.12.21.682, Guernsey. 1967.8.10.2, Scilly Is. 1984.2.26.31, Gulland Rock, 
Padstow, Cornwall. 

ETYMOLOGY. The species was first recognised in material from Guernsey. 

DESCRIPTION. In the erect part of the colony, branching is primarily bifurcate, ranging from equally 
dichotomous, to almost rectilinear series with side branches. The angle between sister stolons 
remains approximately 60 deg. There is a strong disposition towards the rectilinear condition 
where at a bifurcation one daughter stolon tends to remain in line with the main axis of the 
maternal stolon; the other daughter stolon appears sequentially on alternate sides, produced at 
approximately the same distal inclination to the maternal stolon axis as the maternal stolon 
autozooids. The lateral angular inclination of this daughter stolon is about 30 deg. to the orien- 
tation of the maternal autozooid group. Autozooid groups occur at the extreme distal ends of 
stolons, frequently overlying the subsequent branching point. Stolons are usually shaped in 
accommodating the autozooids, being deflected posteriorly at the proximal end of the autozooid 
group. At their distal ends, stolons often broaden, as if to subdivide, providing bases for the 
subsequent daughter stolons, and usually curving anteriorly around the distal end of the autozooid 
group. Occasionally a trifurcation occurs, three daughter stolons being produced. The third stolon 
arises from a posterior projection at the broadened distal end of the maternal stolon; viewed 
anteriorly, this region retains a bilateral symmetry. At the proximal end of the autozooid group, 
autozooids are inclined distally at about 30 deg. to the stolon main axis. The autozooid group 
profile tends to be level at the proximal end of the autozooid group, decreasing at the distal end; this 
is due in part to increasing inclination of the autozooids, and in part to diminishing autozooid 
height. The profile of the rims of the autozooids usually reflects the angle of inclination in having a 
stepped appearance. Viewed anteriorly, a proximal-most autozooid is usually evident in each 
autozooid group. The occurrence of this autozooid, the pattern of autozooid displacements, and 
the succession states of autozooid displacements on the stolons, is identical to that found in A. 
lendigera (see page 313). The orientation of autozooids about the stolonal axis is not rigidly 
preserved from stolon to stolon, with variations up to 90 deg. being possible. Over an area, the sum 
total of such variations is to an extent self cancelling, so that autozooids, overall, face in approxi- 
mately the same direction i.e. in towards a central axis, and thus a relatively sheltered colony- 
bounded space (see page 341). Rhizoids are produced from the anterior face of stolons, just 



AMATHIA 317 

proximal to the autozooid groups. These arise singly, or as a pair, one on either side of the stolon, at 
about 30 deg. to the orientation of the autozooids. 

SI. 1.75-2.75 Z/S.50% 

Sd. 0.80 Zn. 8-23 (appearing as 4- 11 'pairs') 

Zh. 0.38-0.50 

Zw. 0.10 

REMARKS. There is much overlap in the characteristics of A. lendigera, A. intermedis and A. 
guernseii, and it can be very difficult to distinguish among them unless there is an adequate amount 
of material. A.guernseiimay be distinguished on the following basis: the autozooids of A. guernseii 
have a pronounced distal inclination through the entire autozooid group, whereas they tend to 
remain erect in A. intermedis; in A. lendigera, the condition of the autozooids is intermediate. 
A. guernseii is the only species of the three to produce rhizoids. This in turn affects the overall form 
of the colonies; A. lendigera and A, intermedis being diffuse, (the latter also tending to be less 
compact), whereas A. guernseii, with its aggregating rhizoid system, has a more organised and 
directional appearance. These differences would appear to be independent of the type of 
substratum. The description of A. lendigera given by Prenant and Bobin (1956) probably includes 
A. guernseii, as they mention the presence of rhizoids. In all three species, some twist of the stolons 
can occur and this is reflected in the autozooids, but it is never consistent throughout the colony, as 
in A.pruvoti (see pages 336, 337). 

As with A. lendigera, the displacement of the proximal-most autozooids in maternal and 
daughter stolons may reflect the timing of the production of the daughter stolons relative to each 
other (see page 314). 

The holotype is an alcohol-preserved specimen, originally a single colony, now divided into two 
fragments. The substratum is not present in any of the specimens examined. 

DISTRIBUTION. The species is known only from the localities of the type material. 

Amathia populea Busk MS in d'Hondt, 1983 
(Figs 2B, 6D, 7D) 

Amathia lendigera: O'Donoghue & de Watteville, 1944: 430. 

Part Amathia populea Busk MS in d'Hondt, 1983: 97, pi. 3 (4). 

Not part Amathia populea Busk MS in d'Hondt, 1983: 65, ( A. woodsii). 

MATERIAL EXAMINED 

Lectotype (selected here): BMNH; 1899.7.1.526, Natal, S.A., Busk Collection. 

Paralectotypes: BMNH; 1822.8.22.1, Port Alfred, Pondoland, S. Africa. 1851.3.12.36, Port Natal, S. 
Africa 1899.7.1.1 12 C, 513, 540, Algoa Bay, S. Africa. 

OTHER MATERIAL 

BMNH; 1886.7.2.9, 1985.3.4.1, Algoa Bay, S. Africa. 1942.8.6.15, Isipingo Beach, Durban, S. Africa. 
1963.2.14.7, Cape of Good Hope. 

MM; 7061/2, Grahamstown, S. Africa. 7062/2, S. Africa. 7076/2, no locality. 7077/2, Cape Agulhas, S. 
Africa. 

DESCRIPTION. In the erect part of the colony, branching is bifurcate; rarely, a trifurcation occurs. At 
a bifurcation, the two daughter stolons are produced laterally to anterolaterally, at approximately 
30 deg. and 60 deg. to the maternal stolon axis, respectively. The two angular displacements may 
vary, but occur on alternate sides at successive bifurcations. Development in parts of the colony 
may be directionally biased giving rise to plumes of stolons. Plumes may be up to 7 cm. in length, 
with those stolons forming the central axis appearing sympodially arranged. This axis is in fact a 
simple linear series of stolons with lateral branches occurring on alternate sides. Side branches 
within a plume are usually limited to 4 or 5 stolons in sequence. As a result of daughter components 
frequently being produced in a slightly anterior direction, plumes are arc-shaped to cylindrical in 
cross-section. All sequences end with the production of paired lanceolate processes, each process 
made up of 2-3 sequential, progressively tapering kenozooids. Sometimes, the production of a 
stolon in a side branch is replaced by that of a lanceolate process. Autozooid groups reach to the 



318 P. J. CHIMONIDES 

distal ends of stolons, frequently overlying the subsequent branching point. Stolons are usually 
shaped in accommodating the autozooids, appearing raised at the proximal end of the autozooid 
group, becoming shallower distally and usually curving anteriorly to the region of bifurcation. 
Autozooids are outer-wall thickened, but they often appear cylindrical. Autozooid group profile 
diminishes distally, in part due to stolon shape, in part due to decreasing autozooid height. 
Autozooids incline distally at about 30 deg. this being displayed at the autozooid rims, the rims 
usually having a stepped appearance. Viewed anteriorly, a proximal-most autozooid is usually 
evident in each autozooid group. The occurrence of this autozooid, the pattern of autozooid 
displacements, and the succession states of autozooid displacements on the stolons, are identical to 
those found in A. lendigera (see page 313). As one daughter stolon tends to remain in line with its 
maternal stolon, the alternating sequence of autozooid displacements on linear series of stolons is 
more prominent. The orientation of autozooid groups is generally well preserved from stolon to 
stolon. Rhizoids may be produced at the proximal end of stolons, most frequently from those in the 
central axis regions of plumes. Where rhizoids are produced, it is as one per stolon, each arising 
usually from the outer faces of bifurcations, orientated at between 90-180 deg. to the autozooids 
on the same stolon. The resulting colony form is usually arborescent. Secondary development may 
occur in the erect part of the colony where stolons in the common bases of plumes resume normal 
budding of daughter stolons. The angular displacements described above are retained, but without 
maintaining the autozooid orientations about the stolons, or the directional organisation evident 
elsewhere in the colony. The ensuing compact, cotton-wool like, mass may engulf the plume and 
trunk regions. 

SI. 1.00-1.40 Z/S. 30-55% 

Sd. 0.13-0.18 Zn. 6-13 (appearing as 3-6 'pairs') 

Zh. 0.38 Tpl. 1.75 (2.60 max.) 

Zw. 0.10-0.13 

REMARKS. An association with a sandy environment is inferred from the sand grains sometimes 
found accreted to rhizoids and attached epizoic worm tubes. In plume portions of the colony, the 
preserved orientation of the autozooid groups, and the cross-sectional profile of the regions, results 
in autozooids facing into a relatively sheltered colony-bounded space (see page 341). 

The plume portions of A.populea strongly resemble the figures of A. lemaniiin the unpublished 
plates of Lesueur. However, it is equally possible to draw a similarity between these figures and A. 
woodsii (see page 324) or possibly portions of A. tricornis. 

Understandably, A.populea has, in the past, been confused with A. woodsii and A. tricornis (e.g. 
d'Hondt, 1979, 1983). It has also been confused with A. lendigera (e.g. O'Donoghue and de 
Watteville, 1944, BMNH 1942.8.6.15). A. populea may be distinguished from A. lendigera (and 
similar forms A. guernseii and A. intermedis) primarily through the occurrence and location of 
rhizoids. These do not occur in A. lendigera or A. intermedis. In A. guernseii, the rhizoids are 
produced anteriorly, just proximal to the autozooid group; whereas in A. populea they are pro- 
duced latero-posteriorly and proximally distant from the autozooid group. A. tricornis and A. 
populea differ in many characteristics (see page 321). 

D'Hondt (1979) placed A.populea Busk MS (part, without qualification) into synonymy with A. 
cornuta (sensu d'Hondt, 1979) along with a number of other species, including A. australis. 
D'Hondt (1983) then drew some distinction, first indicating (p. 65) that A. populea Busk MS part 
from Australia is synonymous with A. cornuta (sensu d'Hondt, 1983 i.e. A. woodsii see pages 320, 
323 etseq.}. Later, d'Hondt (1983: p.97) also gave a brief description and a figure (p. 103) of a South 
African specimen, BMNH 1899.7.1.526 of A. populea Busk MS part, so validating Busk's manu- 
script name, and making the name A.populea available for this species. D'Hondt referred to the 
specimen as 'A.sp.' yet appears to have remained equivocal by suggesting that this is also possibly 
'a form of A. cornutaT (sensu d'Hondt, 1983) i.e. A. woodsii (see pages 320, 323 et seq.). 

A. populea and A. woodsii may be distinguished in the following: the form of the lanceolate 
processes, being simple in A. populea, often branched in A. woodsii; the autozooid to stolon ratio, 
being higher in A. woodsii; the orientation of the rhizoid origins, being latero-posterior in A. 
populea and anterior in A. woodsii. 



AMATHIA 319 

Small quantities of material may be very difficult to distinguish and identify with certainty, such 
that even Busk made errors. Some of Busk's A. populea, BMNH 1899.7.1.528 from Algoa Bay, 
South Africa and BMNH 1899.7.1.4383 from Australia, is in fact A. woodsii. BMNH 
1899.7.1.4383 is the only specimen in the BMNH collections from Australia labelled A. populea, 
and so is undoubtedly the material that d'Hondt (1983) refers to under the name "A. populea Busk, 
unpublished (pars: Australia)'. 

All specimens labelled by Busk as A. populea and considered by d'Hondt (1983) are certain 
syntype material. D'Hondt's figured specimen, (BMNH 1899.7.1.526) is here chosen as lectotype, 
the remaining Busk material, except for the two misidentifications indicated above, has 
paralectotype status. 

DISTRIBUTION. The species is known from the south-eastern coast of South Africa, possibly also 
occurring off southern Australia. 

A mat Ida woodsii Goldstein, 1879 
(Figs 2C, 9B, D) 

Amathia woodsii Goldstein, 1879: 20, pi. 3 (fig. 5). 
Amathia australis: MacGillivray, 1889: 310, pi. 185 (figs 5, 5a). 
Amathia woodsii: MacGillivray, 1895: 138, pi. B (figs 5, 5a). 
Part Amathia cornuta: d'Hondt, 1983: 65, fig. 36 (C). 

MATERIAL EXAMINED 
Neotype (selected here): BMNH; 1883.1 1.29.27, Port Jackson. 

OTHER MATERIAL 

BMNH; 1861.9.20.17, Fremantle. 1897.5.1.1189, no locality. 1897.5.1.1196, Port Phillip Heads. 
1899.7.1.528, 1985.3.6.1, Algoa Bay, S. Africa. 1899.7.1.4383, Australia. 1909.8.4.10, Western Port, 
Australia. 1963.3.28.4, Adelaide. 

MM; 7075/2, Queensland. 

DESCRIPTION. In the erect part of the colony, branching is bifurcate. Stolons are arranged to form 
rectilinear series with side branch stolons. Side branch stolons are produced on alternate sides at 
each subsequent bifurcation, arising with the same distal inclination as the autozooids of their 
maternal stolons. The lateral angular displacement of the side branch stolons can be 0-90 deg. to 
the autozooid orientation, but usually ranges from 10-30 deg. In parts, growth appears favoured 
along the rectilinear series, with side branches usually restricted to 4-5 stolon units either side. 
These parts of the colony have a plume like appearance. Branches end with a pair of lanceolate 
terminal processes, usually produced in the same orientations as stolons. The processes are made 
up of 3-4 sequential, progressively tapering kenozooids, often bifurcating at the distal end of the 
basal segment kenozooid. The lanceolate processes in which bifurcation occurs are most usually 
produced in the non-rectilinear position. Frequently, the production of a side branch is replaced by 
the production of a lanceolate process, emphasising the appearance of directional growth. Auto- 
zooid groups occur towards the distal ends of stolons, but often there is further autozooid-free 
part, coinciding with the production of a side branch component. Stolons may show a gentle 
anterior curvature, and sometimes curve around the distal autozooids of a group. Autozooid 
group profile diminishes distally, mainly due to decreasing autozooid height, but sometimes due in 
part to an increase in their distal inclination. Autozooids are outer-wall thickened, the walls 
appearing cylindrical. Viewed anteriorly, a proximal-most autozooid is usually evident in each 
autozooid group. The occurrence of this autozooid, the pattern of autozooid displacements, and 
the succession states of autozooid displacements on the stolons, are identical to those found in A, 
populea (see page 318). The orientation of autozooid groups is generally well preserved from stolon 
to stolon. Along a plume therefore, autozooids on the rectilinear sequence all face in the same 
direction, with lateral stolon autozooids generally facing across these. Rhizoids may be produced, 
one per stolon, arising near to and at about the same orientation as the autozooids. Colony 
arrangement is similar to A. populea. 



320 P. J. CHIMONIDES 

Zh. 0.65 Z/S. 50-70% 

Zw. 0.10 Zn. 8-23 (appearing as 4- 11 'pairs') 

SI. 2.58 Tpl. 2.50 (4.80 max.) 

Sd. 0.20 

REMARKS. Goldstein's account and figure are a very good representation of the species; the only 
omission is information on rhizoid production. D'Hondt (1983) placed the species in synonymy 
with A . cornuta Lamarck (1816), but there is some doubt as to the identity proposed for Lamarck's 
specimen by d'Hondt (1983), and the distinction between A. cornuta Lamarck and A. woodsii is 
here maintained (see page 323 et seq.}. 

With limited material, confusion could arise between A. woodsii and A.populea or A. tricornis. 
D'Hondt (1983) has referred specimens of these last two species to A. cornuta Lamarck (sensu 
d'Hondt 1983), i.e. A. woodsii. The species may be distinguished in the following: the presence of 
the characteristic subdivided lanceolate process in A. woodsii, this being simple in the other two; the 
production of rhizoids being near to, and in the same orientation as the autozooid group in A. 
woodsii, these being distant, and of different orientation in the other two; the budding pattern in A. 
woodsii is never as complex as in A. tricornis, and the predisposition to rectilinear development is 
more prominent than in A.populea, in which there is a tendency for a sympodial appearance. 

According to Stach (1936) specimens from Goldstein's collection were deposited in the NMV. 
However, his material for A. woodsii is not there (NMV in litt. 6.12.1984). In view of the confusion 
which has arisen, there is a need for type material. The description and measurements given here 
are based on BMNH 1883. 1 1 .29.27 from Port Jackson, an alcohol specimen, rehydrated from the 
dry state. The specimen is here selected as neotype. Goldstein does not give a locality for his 
specimen, only mentioning that the species was found on a previous occasion at Portland, 
presumably Victoria State. 

DISTRIBUTION. The species is known from the south-eastern coast of South Africa, and from 
Australia, with records from Fremantle, Adelaide, Port Phillip Heads, Port Jackson and 
'Queensland'. 



Amathia tricornis Busk MS 
(Figs2D, 12C) 

Holotype: BMNH; 1899.7.1.6600, Australia, Busk Collection. 
Paratypes: BMNH 1899.7.1.4393, 4394, Australia, Busk Collection. 

ETYMOLOGY. Busk's MS name, probably indicating the occurrence of three terminal lanceolate 
processes. 

DESCRIPTION. In the erect part of the colony, branching is mainly trifurcate, although tetrafur- 
cation occurs in certain regions. The latter condition is associated with astogenetically early parts 
of the colony, which form the base and central main-axis regions. These regions tend to be 
composed of lengthy series of, what are here termed, type 'a' stolons. Type 'a' stolons are longer 
than other type 'b' stolons found in the trifurcate portions of the colony, and often bear rhizoids. 
Where rhizoids are produced, these arise from the proximal end of a stolon, usually singly, at 
between 45-90 deg. to the autozooid orientation on the same stolon. Autozooids borne by type 'a' 
stolons show no difference in size from those on type 'b' stolons, although the autozooid groups 
tend to be shorter. The linear proportion of stolon occupied by autozooids, therefore, is lower. The 
stolon budding arrangement, in both tri- and tetrafurcate conditions, always results in one 
daughter stolon lying in rectilinear succession to the maternal stolon. Two other daughter stolons 
are produced laterally, one on each side, at about 60 deg. to this central axis. All three of these 
daughter stolons bear autozooids, usually orientated in the same direction as those on the maternal 
stolon, with some exceptions. In the tetrafurcate condition, a fourth daughter stolon is produced, 
also at about 60 deg. to the central axis, but posteriorly to the maternal stolon. The autozooid 
group orientation of the maternal stolon is preserved in this daughter component; the autozooid 



AMATHIA 321 

group thus faces distally along the rectilinear series of the central axis (see Fig. 2D). In the 
tetrafurcate condition alone, the orientation of the autozooid groups along the rectilinear series is 
not always maintained. A repeat rotation of 90 deg. may, instead, be observed at each axial 
junction. The relationship of sister daughter stolons to the axial daughter stolon remains fixed, 
and, thus, the entire assemblage follows the re-orientation. The original orientations are recovered 
every fourth axial stolon unit along the sequence. In the trifurcate condition, lateral growth 
appears to be limited to one or two stolons in sequence each side. Development in these parts of the 
colony is therefore directionally biased and these parts have a plume like appearance. Branches end 
with the production of three lanceolate terminal processes, each made up of two or three sequen- 
tial, progressively tapering kenozooids. These arise from, and lie approximately in line with, their 
maternal stolons. Sometimes, the central terminal process does not form, being replaced by a 
stolon instead. This may be repeated so that occasionally, lateral branches may be several stolons 
in length. On both type 'a' and type 'b' stolons, autozooid groups occur at the extreme distal end of 
stolons, frequently overlying the subsequent branching point. Autozooid group profile diminishes 
distally in all cases, mainly due to increasing distal inclination of the autozooids. In all parts of the 
colony, autozooids are outer-wall thickened. Along rectilinear sequences of stolons, there is a 
predictable repetition in the arrangement of the autozooids borne. The sequence, progressing 
distally, is as follows: if, in an autozooid group, there is one proximal-most autozooid prominent, 
this is associated with one side of the stolon; in the next stolon, no single autozooid is prominent 
proximally, the proximal autozooids being paired equally; on the third stolon, a proximal-most 
autozooid is prominent once again, but on the opposite side to that of the first stolon; on the fourth 
stolon, the proximal autozooids are paired as on the second stolon; the fifth stolon repeats the 
arrangement on the first stolon. On laterally branched stolons, a proximal-most autozooid is 
prominent, and is associated with the side nearest the rectilinear stolon sequence. Autozooids on 
stolons continuing in rectilinear series which develop from lateral branch stolons subsequently 
follow the predictable pattern of repetition given above. 

SI. (a) 2. 10 Z/S. (a) 30-40% 

Sd. (a) 0.25 Zn. (a) 8-1 1 (appearing as 4-5 'pairs') 

SI. (b)1.45 Z/S. (b) 60% 

Sd. (b)0.25 Zn. (b) 10-21 (appearing as 5-9 'pairs') 

Zh. 0.35 (all autozooids) 

Zw. 0.10 (all autozooids) 

Tpl. 2.10 

REMARKS. No evidence exists, in the limited material available, that the trifurcate condition ever 
gives rise to the tetrafurcate condition. The colony form is inferred to be arborescent, resulting 
from the production of rhizoids. 

None of the material held at the BMNH named A. tricornis in MS by Busk is misidentified. 
D'Hondt (1979 & 1983) erroneously placed this species in synonymy in part, with parts of A. 
cornuta (Lamarck) sensu d'Hondt (i.e. A. woodsii, see page 323), and in part, initially with A. 
pinnata (1979) (see page 330), and subsequently with A. inarmata (1983) (i.e. A. biseriata, see page 
332). D'Hondt on each occasion mentioned A. tricornis in synonymy only, thus not making the 
name available at any time (I.C.Z.N. article 1 le). 

Among the species which may be confused with A. tricornis are: A. populea; A. woodsii; A. 
pinnata; A. biseriata. In brief, A. tricornis has a more complex colony construction and differs from 
these species in many features, for example: the bimorphic autozooid-bearing stolons; the normal 
occurrence of tetra- and trifurcation, including the production of triplet lanceolate processes and 
their permutations with autozooid-bearing stolons; autozooid and stolon re-orientations; the 
productions site of the rhizoids. There are also differences in the autozooid to stolon ratios. 

DISTRIBUTION. The species is known only from material described as being from 'Australia', sent to 
Busk by Miss Gore. 



322 P. J. CHIMONIDES 

Amathia lamourouxi nom. nov. for 

Amathia cornuta auctorem 

(Figs 3A, 8C, 9A, C) 

? Not Serialaria cornuta Lamarck, 1816: 131. 
Amathia cornuta Lamouroux, 1816: 159, pi. 4 (fig. la, IB). 
? Not Serialaria australis Tenison Woods, 1877: 83, 1st fig. 
? Not Amathia australis: Tenison Woods, 1880: 102. 
Amathia cornuta: Tenison Woods, 1880: 99, fig. 3. 
Not Amathia australis: MacGillivray, 1889: 310, pi. 185 

(figs 5, 5a), ( = A. woodsii). 

Amathia cornuta: MacGillivray, 1895: 137, pi. D (fig. 1, la). 
? Amathia cornuta: d'Hondt, 1979: 10, 16. 
Part Amathia australis: d'Hondt, 1983: 65, fig. 36(F). 
Not Amathia cornuta: d'Hondt, 1983: 65, fig. 36(C) ( = A. woodsii). 

MATERIAL EXAMINED 
Neotype (selected here): BMNH; 1887.12.10.70, Port Phillip, J. B. Wilson collection. 

OTHER MATERIAL 

BMNH; 1842. 11. 4.50, Sydney. 1899.7. 1.3, New Zealand. 1899.7.1.4325, Victoria. 1899.7.1.4327,4329-31, 
4333, Australia. 1899.7.1.4328, Bass Strait. 1899.7.1.4334, Australia & New Zealand. 1985.3.10.1, Flinders 
Is., Bass Strait. 1985.3.24.1, mid channel, Port Phillip Heads, 15m. 

MM; 7078/2, Australia. 

LBIMM; bry 2821 part, Australie Occidentale/Nouvelle Hollande (see below). 

ETYMOLOGY. Lamouroux's name is used for his species of A . cornuta ( 1 8 1 6), a name preoccupied by 
A. cornuta (Lamarck, 1816). 

DESCRIPTION. In the erect part of the colony, branching is bifurcate. At any branching point, 
daughter stolons may be produced in positions ranging from almost rectilinear, to 90 deg. to the 
maternal stolon. A minimum separation of 90 deg. occurs between daughter stolons. These usually 
arise from the posterior side of their maternal stolon. Stolons are narrowed proximally and usually 
curved anteriorly, being reminiscent of a short, simple, cow horn. Autozooid groups occupy the 
greater part of stolons, and frequently overlie the subsequent branching point. Autozooid group 
profile usually increases proximodistally within each group, or may remain level. Autozooids are 
outer-wall thickened. Viewed anteriorly, a single proximal-most autozooid is evident in each 
autozooid group. This autozooid is usually placed just off the stolon mid-line, thus associated with 
either one or other side of the autozoid group. No pattern is evident from group to group, in the 
location of this autozooid. At the distal end of each autozooid group and contiguous with the 
autozooids, are produced a pair of tapering, single-kenozooid lanceolate processes. At the distal 
extremities of the colony, both daughter stolons tend to be produced at 90 deg. to their maternal 
stolons, i.e. separated from each other in equal dichotomy by 180 deg. In less distal parts of the 
colony, daughter stolons may be separated from each other in equal dichotomy by lesser angles. 
Daughter stolons produced in the linear position are less common and are associated with more 
central and proximal (astogenetically earlier) regions of the colony. Autozooid orientation is 
generally not preserved from stolon to stolon. Often there is an equal rotation by up to 90 deg. of 
each daughter stolon in opposite directions i.e. were both daughter stolons to lie in the linear 
position, their anterior faces would be away from each other. Successive daughter stolons actually 
lying in the linear position and forming a sequence, are all produced from the same side, i.e. in such 
a sequence, viewed anteriorly each time, they would all have been budded from e.g. the left side. 
The orientation of the autozooid groups, in such a linear sequence, is rotated by 90 deg. in the same 
direction, with each successive stolon. The original orientation is recovered every fourth stolon 
unit. Superficially, branching can appear as 'alternate' along a linear sequence. Rhizoids may be 
produced, usually from stolons along these linear sequences. Rhizoids are produced one per 
stolon, arising from a position level with, and at 90 deg. to the proximal-most autozoid, and on the 
same side of the stolon as the direction in which the stolon was budded. 



AMATHIA 323 

Zh. 0.40-0.50 Z/S.75% 

Zw. 0. 1 3 Zn. 1 1-1 5 (appearing as 5-7 'pairs') 

SI. 1.13 Tpl. 2.00-2.25 

Sw. 0.25 (at the widest region) 

REMARKS. No pattern has been discerned in the autozooid arrangement from stolon to stolon, 
other than, if the proximal-most autozooid of a group is associated with one side of the stolon, then 
that association may remain in both daughter stolons over a number of successive bifurcations. 

The identity of A. cornuta auct. is inextricably associated with the collections made by Peron and 
Leseur, between the years 1800-1804 (see page 307), on which both Lamarck and Lamouroux 
worked, both of them describing a 'cornuta'. 

Tenison Woods (1880) drew a distinction between his A. australis and A. cornuta sensu 
Lamouroux (i.e. A. lamourouxi) based on the understanding that Lamouroux's (1816) figure 
indicates a single line of autozooids. Lamouroux himself, referred in the singular to 'the largest cell 
of each group . . . garnished with two setaceous appendages'. The misinterpretation of a single row 
for a double row of autozooids, might be made as a result of a preservation artifact where, in dried 
specimens, the thinner central walls between autozooids collapse from view, leaving only the outer 
walls visible. On this basis there is no distinction between A. australis and A. lamourouxi. The 
additional difference claimed by Tenison Woods, in the form of the 'setaceous appendages' (his 
figure of 1 877 shows these as being broad and less trim than those in Lamouroux's figure), might be 
accounted for in terms of the variation which may occur within A. lamourouxi. However, Tenison 
Woods' figure (1877) shows clearly that his specimen had undergone trifurcation. Two possibilities 
may account for this: the first, that under certain conditions, A. lamourouxi can undergo such a 
division; the second, that Tenison Woods did in fact have a separate species. Although the former 
may be possible, trifurcation has not been recognised in specimens here assigned to A. lamourouxi. 
The whereabouts of Tenison Woods' material is not known. 

MacGillivray (1889) considered A. australis to be A. cornuta sensu Lamouroux, but in his 
description (p. 310) and figure (pi. 185, figs 5, 5a) gave an acocunt of A. woodsii. MacGillivray 
(1895) subsequently recognised the error, and correctly referred to his account of 1889 as being 
descriptive of A. woodsii. At the same time, MacGillivray distinguished between A. woodsii and A. 
cornuta, and reaffirmed his opinion that A. australis was synonymous with the latter, but gave 
Lamouroux as the author and placing Lamarck in synonymy. 

D'Hondt ( 1 979) has found a specimen, LBIMM bry 282 1 , which is claimed to be the holotype of 
A. cornuta (Lamarck). D'Hondt (1979) placed A. australis in synonymy under A. cornuta 
(Lamarck), but without mention of A. woodsii. D'Hondt (1983) then placed A. woodsii in 
synonymy under A. cornuta (Lamarck), but excluded A. australis, thus revoking his opinion of 
1 979 and indicating that A . australis is different (d'Hondt's reference to 'parts' of A. australis at this 
point are enigmatic). In this reorganisation of the species, d'Hondt (1983) gave two figures: 36(C) 
as A. cornuta (Lamarck), and 36(F) as A. australis. Figure 36(C) is in fact A. woodsii, and 36(F) is A. 
cornuta, both of common usage, the latter corresponding with Lamouroux (1816), of which A. 
australis is usually taken to be a junior synonym. 

D'Hondt (1983) appears to have determined A. cornuta (Lamarck) to be different from A. 
cornuta Lamouroux. A. cornuta (Lamarck) predates A. cornuta Lamouroux (Tenison Woods, 
1880, d'Hondt, 1983). D'Hondt (1983) thus relegated A. woodsii as a junior synonym of A. cornuta 
(Lamarck), and assigned the name A. australis, as the next available name, for what was previously 
accepted as A. cornuta sensu Lamouroux. 

Unless it was the only specimen involved, LBIMM bry 2821 can only be taken as the holotype if 
so designated at the time of introduction by the original author. Lamarck (1816) did not do this, 
and the number of specimens involved is not certain. 

LBIMM bry 2821 is recorded as being one of three specimens of A. cornuta so identified by 
Lamarck and in the Paris Museum at the time of the compilation of the first catalogue of Bryozoa 
in 1 867. The other two specimens appear to have been A . cornuta sensu Lamouroux, only 'possibly' 
originating from Peron and Lesueur. Their locality is given as 'Australasie'. The specimens were 



324 P. J. CHIMONIDES 

numbered: 172a,b,c. (photocopy of the 1867 catalogue). LBIMM bry 2821 is the only one of the 
three which is known to have come from Peron and Lesueur (d'Hondt in litt. 10.12.1984). The 
locality for this specimen is 'Australie Occidental' (d'Hondt, 1979), and also as 'Nouvelle- 
Holland' (loan form 24th Jan. 1985). 

D'Hondt (1979) reported that LBIMM bry 2821 carries the label 'Amathia lemanii Lesueur'. 
This would appear to be in the hand of Pergens, the original label having been lost or destroyed. 
The specimen is taken to be the same one that Pergens (1887) correlated with a figure in the 
unpublished plates of Lesueur, these in turn related to a manuscript of Desmarest and Lesueur, 
deposited at Paris in 1 829 (with another slightly different version at le Havre). Pergens ( 1 887, p. 88) 
ascertained that plate 13, figure 6, in the unpublished plates, is Amathia lemanii, and (p. 90) then 
gave the identification he was able to make of the species in terms of what was, to him, a valid and 
available name i.e. A. cornuta (Lamarck). Copies of the plates of Lesueur exist at the BMNH. 
Plate 13, figure 6, consists of three representations of the species intended, all three at different 
magnifications. The species represented could be A. woodsii or A. populea (see page 318); both 
species are capable of assuming the characteristics portrayed. Missing from the figure(s) is any 
information on rhizoids and on any occurrence of the characteristic subdivided terminal filaments, 
which might serve to distinguish between the two species. Only the actual specimen used by 
Lesueur will determine the true species (taken to be LBIMM bry 2821). The identity of this species 
is of little taxonomic consequence however, as neither plates nor descriptions have ever been 
published. LBIMM bry 2821 could have been the holotype perhaps, of Desmarest and Lesueur's 
species, but there is insufficient evidence published to suggest that it was that of Larmarck's. 

In addition, Larmarck (1816) gave no figure, specimen number, or dimensions, with which a 
specimen may be correlated. Furthermore, the locality information (see above) for the specimen, 
although close, does not match exactly with that of Lamarck (or of Lamouroux). Lamarck gives 
TOcean asiatique' (Lamouroux gives 'Sur les Fucus de 1'Australasie'). That the specimen was part 
of Peron and Lesueur's collections, may not in this case be sufficient; Lamarck himself is not 
definite as to the origins of his specimen, only 'believing' it to be from Peron and Lesueur. Pergens 
(1887) merely expressed his opinion that LBIMM bry 2821 is the same as A. cornuta (Lamarck); 
how he reached that conclusion is not clear. The specimen appears to be only one remaining of a 
number, others having gone astray since the days of Peron and Lesueur; the 1867 catalogue of the 
Bryozoa was compiled some 51 years after Lamarck's publication. 

It is possible that LBIMM bry 2821 may be eligible for selection as lectotype of A. cornuta 
(Lamarck), if it can be shown to have been part of Lamarck's original syntype series, and formative 
of his opinion. However, Larmarck did also identify two different specimens as being his species, 
these being A . cornuta sensu Lamouroux (see above) . These specimens might also have been eligible 
for selection, but are no longer to be found in the Paris Museum (d'Hondt in litt. 10.12.1984, 
24.01.1985). 

Further challenge to the identity proposed by d'Hondt for A. cornuta lies in the evidence 
that preceeds Pergen's opinion (1887). Tenison Woods (1880) gave information on the working 
relationship between Lamarck and Lamouroux concerning the Amathia specimens collected by 
Peron and Lesueur. Much of the information appears to be derived from Lamouroux's (1816) own 
preface and introduction. Lamouroux had 'the fullest access' to Lamarck's collection, and named 
at least part of this, if not all of it. 

Neither Lamarck's (1816) nor Lamouroux's (1816) account of a 'cornuta' contradicts the other. 
However, whereas Lamouroux's account is quite explicit, and furnished with figures, such that the 
species he described may still be recognised; Lamarck's account is open to interpretation. The 
descriptions may be translated as follows: 

Lamouroux, p. 159: No. 266. 

(from the French) The largest cell of each group having its free border, garnished with two 

setaceous appendages. 

(from the Latin) two setaceous filaments from the first rank cell 

Lamarck, p. 131: No. 2. 

(from the French) I believe it (to be) from the voyage of Messieurs le Sueur and Peron. It is a 



AMATHIA 325 

little more stout and less capillary than the preceeding [i.e. A. lendigera], at the extremities curved 
and as curls. 

(from the Latin) very branched, articulated, somewhat curled; branches alternate; curved 
secondary little branches; cells in distinct series; two setae at the most distant extremity. 

In Lamarck's account, no orientation is given for the setae, and the description could apply to A. 
cornuta of Lamouroux, A . woodsii or A . populea. Whether the reference to the secondary branches 
is an indication of an arborescent growth form i.e. axial development with lateral branch system, or 
a reference to the stolons themselves, is not clear. In either case, the description is insufficiently 
distinctive. Branching is alternate in A. woodsii and A. populea and may appear so in A. cornuta 
sensu Lamouroux. Finally, Lamarck makes no mention of any subdivided terminal processes 
(present in LBIMM bry 2821 part) to be expected if his 'cornuta' was the equivalent of A. woodsii. 

Although it is not possible to recognise a single species from Lamarck's description, the identity 
of A. cornuta (Lamarck) has been understood through the later accounts of the two authors: in 
Lamouroux (1824) and Lamarck (1836) respectively, each recognises the other's A. cornuta as 
synonymous with his own; from this derives the A. cornuta of common usage. It is this concordance 
which d'Hondt (1983) has in effect repudiated. 

In the strictest sense, A. cornuta (Lamarck) should have been classed as a nomen dubium, and not 
used. This is historically implied by Tenison Woods (1880) who acknowledged that Lamarck 
probably predated Lamouroux, and so accepted A. cornuta (Lamarck), 'but with reference to 
Lamouroux only'. MacGillivray (1895) appears to have been of the same opinion (see above). 

In the light of such contradictions, Lamarck's A. cornuta must be taken as a nomen dubium, and 
the name should no longer be used for Lamouroux's species. A. australis of Tenison Woods would 
be the next valid name available, if certainty could be attached to the identity of his species (see 
above). Under these circumstances, it is wiser, in the interest of long term stability, to select a new 
name for A. cornuta Lamouroux, accepting either his figures as lectotype, or perhaps selecting a 
neotype. It is here proposed that A. cornuta auct. be known as A. lamourouxi, with specimen 
BMNH 1887.12.10.70 as neotype. 

DISTRIBUTION. The species is recorded from New Zealand and southern Australia. 

Amathia plumosa MacGillivray, 1890 
(Figs3C, 12A, B) 

Amathia plumosa MacGillivray, 1890: 110. 

Amathia plumosa: MacGillivray, 1895: 139, pi. C (figs 2, 2a). 

Amathia plumosa: d'Hondt, 1983: 67, fig. 36 (B). 

MATERIAL EXAMINED 
Holotype: NMV; H494, Port Phillip Heads, J. B. Wilson Collection. 

OTHER MATERIAL 

BMNH; 1963.2.12.354, 358, Western Australia. 1985.3.8.1, no locality. 

DESCRIPTION. The branching pattern on the erect part of the colony is based on both bi- and 
trifurcation. Trifurcation is associated with non autozooid-bearing stolonal kenozooids, forming 
angularly undulating 'main-stems'. At the distal end of each of the main-stem constituent stolonal 
kenozooids, are produced: a single continuing stolonal kenozooid, deflected by approximately 30 
deg. towards the central axis of the main-stem; two (autozooid-bearing) side branch stolons, one 
each side. The side branch stolons are produced in the same plane as their maternal stolonal 
kenozooid, but diverge from each other equally, by an approximate total angle of 60 deg. The 
autozooids borne on these side branch stolons face the main-stem, and the stolons themselves are 
curved anteriorly. Subsequent branching from these side branch stolons is usually bifurcate, 
although new main-stem sequences may be produced, showing the associated trifurcation. 
Development along side branches is usually limited; 2-3 stolons in a sequence is usual, but up to 8 
stolons in succession may occur. The orientation of autozooid groups along any such sequence 
remains the same. These side branches end with the production of a pair of usually dichotomously 
branched lanceolate processes. These are made up of sequential, progressively tapering 



326 P. J. CHIMONIDES 

kenozooids. Where the processes are branched, this occurs at the distal end of the base segment 
kenozooid. This may be repeated in one or both of the next resultant segments. Rarely, a lanceolate 
process may trifurcate. On occasions, the production of a stolon in a side branch is replaced by the 
production of a lanceolate process. Development in such cases, therefore, tends to be directionally 
biased. The colony is composed of such quasi-cylindrical assemblages, circular in cross-sectional 
profile, and somewhat reminiscent of 'feather boas'. These may be supported on a trunk-like part 
of the colony, resulting from the production of rhizoids (see page 309). Rhizoids are produced in 
two ways: they may be produced from the proximal end of main-stem kenozooids, sometimes 
singly, although more often as an adjacent pair, in the same orientation as the side branch stolons 
lying immediately proximal; they may be produced from autozooid-bearing stolons, at approxi- 
mately 120 deg. to the orientation of the autozooid group on the same stolons. Where autozooid 
groups develop in side branches, these occur towards the distal ends of stolons, but often there is a 
further autozooid-free portion. This portion is about the same length as the diameter of, and 
coincident with the production of, a daughter component. Autozooid group profile tends to 
remain level and autozooids are outer- wall thickened. Where a proximal-most autozooid of a 
group is evident, its occurrence, and the pattern of autozooid displacements from stolon to stolon, 
are similar to those of A. lendigera (see page 313) except that the second succession state-does not 
appear to occur. 

kSl. 1-45 (main-stem kenozooidal stolons) 

kSd. 0- 1 9-0-29 (main-stem kenozooidal stolons) 

SI. 1-03-1-61 Z/S. 55-65% 

Sd. 0-15-0-26 Zn. 7-17 (appearing as 3-8 'pairs') 

Zh. 0-32 

Zw. 0-10 

REMARKS. The species is so distinctive that it does not appear to have been confused with any 
other. The slide mounted specimen NMV H494, is accepted here as the holotype of A. plumosa 
MacGillivray (1890), and as that figured by MacGillivray (1895). The label on the slide carries the 
information: 'H494 Amathia plumosa McG P.P.H. fig J.B.W.'. This in agreement with the original 
description in which the locality is given as 'Port Phillip Heads', from the collection of J. B. Wilson. 
The description given here is based on the above specimen. Some supplementary information is 
derived from BMNH 1963.2.12.354 and BMNH 1963.2.12.358, these agreeing well with the 
holotype. 

There is some indication that the repeated branching in the lanceolate processes coincides with 
the development of a lanceolate process in substitution for an expected stolon, although there is no 
certainty to this. Autozooid groups are orientated about main-axis stolons to face into relatively 
sheltered, colony-bounded space (see page 341). 

DISTRIBUTION. The species is known only from Australia, recorded from 'western' Australia and 
the type locality of Port Phillip Heads in the south-east. 

Amathia obliqua MacGillivray, 1985 
(Figs 3D, 8B) 

Amathia obliqua MacGillivray, 1895: 135, pi. B (figs 2, 2a). 

MATERIAL EXAMINED 

Syntypes: NMV; H493 (old number 65391), H493 (old number 65392), Port Phillip heads, J. B. Wilson 
Collection. 

OTHER MATERIAL 

MM; 7108/2W, Port Phillip. 

DESCRIPTION. In the erect part of the colony, branching is bifurcate with repetitive gradual varia- 
tion evident in the branching angle. This ranges from equal dichotomy, to the condition where the 
daughter stolons are produced at angles to the maternal stolon axis of 30 deg. and 60 deg. 
respectively. This variation occurs over a sequence of four stolon units, i.e. if one daughter stolon is 



AMATHIA 327 

angled at 60 deg. to the right of a maternal stolon, the same angular displacement appears, to the 
left of a maternal stolon, four stolon units further on in a stolon sequence. The original angular 
displacements are recovered after a further sequence of four stolon units. In between each of these 
stages, there is an intermediate, equally dichotomous condition. Over the entire sequence, a sig- 
moidal pattern in stolon arrangement may be observed. Autozooid groups occur towards the distal 
ends of stolons, but usually there is a further autozooid-free portion beyond the group, of variable 
length. Stolons tend to be straight, but sometimes the distal autozooid-free portion may be twisted 
slightly or deflected anteriorly, or both. Autozooid groups are set obliquely on the stolons. The 
autozooid group points in the same direction as that, in which the bearing stolon itself was budded 
i.e. viewed anteriorly, on a right hand daughter stolon, the autozooid group starts proximally on 
the left of the stolon and finishes distally on the right, and vice versa. Autozooid group orientation 
is generally well preserved from stolon to stolon, though variations of up to 30 deg. may occur. 
Autozooids are outer-wall thickened, and the autozooids of any one group tend to be about the 
same height. Autozooid group profile therefore tends to be level. A single proximal-most auto- 
zooid is usually prominent in each autozooid group. Its occurrence, and the pattern of autozooid 
displacements from stolon to stolon, are similar to those of A. lendigera (see page 313) except that 
the second succession state does not appear to occur. Rhizoids may be produced, one per stolon, 
from mid-way along the proximal autozooid-free end. These are orientated at about 90 deg. to the 
autozooids, on the outer faces of stolons at a bifurcation i.e. on the side of a stolon away from its 
sister stolon. 

Zh. 0-39 Z/S.65% 

Zw. 0-11 Zn. 11-21 (appearing as 5-10 'pairs'). 

SI. 1-16-2-13 

Sd. 0- 1 3-0-20 (immediately proximal to the autozooids) 

REMARKS. Little material is available for study, therefore little is known of the colony form, other 
than from MacGillivray's original description. It is inferred, from the presence of rhizoids, that the 
colony attains an arborescent form. MacGillivray's (1895) description seems to bear this out, the 
colony being 'attached by the bases of main stems by radical tubes, the branches being quite free 
and not intertwining or climbing over other objects'. MacGillivray made no mention of the non- 
erect part of the colony. Neither of the two slide specimens from the NMV, H493 (65391, 65392) 
matches the figure of MacGillivray (1895) exactly. There is however a very close resemblance to 
specimen H493 (65391). Some of this colony fragment has broken away which may account for the 
lack of congruence with the figure. 

As with many other species of Amathia, A. obliqua has been confused with A. lendigera (by 
MacGillivray 1895). The presence of rhizoids and their orientation, the development pattern of 
autozooid groups and the overall colony form, serve to distinguish this species from A. lendigera 
(and also from A. inter medis and A. guernseii). 

DISTRIBUTION. The species is recorded only from the Port Phillip Bay region in Australia. 

Amathia wilsoni Kirkpatrick 1888 
(Figs 4D, IOC, D) 

Amathia wilsoni Kirkpatrick, 1888: 18, pi. 2 (figs 4, 4a). 
Amathia wilsoni: MacGillivray, 1895: 139, pi. D (figs 2, 2a, 2b). 
Amathia wilsoni: d'Hondt, 1983: 67, fig. 36 (A). 

MATERIAL EXAMINED 
Syntype: BMNH; 1888.5.17.7, Port Phillip, J. B. Wilson Collection. 

OTHER MATERIAL 

BMNH; 1821.5.24.16, 1985.3.12.2, Portland, Australia. 1882.7.7.54, Wilsons Promontory. 1886.6.8.3, 
Port Phillip 1910.10.17.31-32 part, north end Victoria Tasman Cable, <50fthms. (91.44m). 
1963.2.12.361, Australia 1963.2.12.366, Holdfast Bay nr. Adelaide. 1985.3. 12.1a,b, Flinders Is. 1985.3.18.3, 
Hobart, Tasmania. 

MM; 7136/3W, off Shark Is., Port Jackson. 7137/3W, Port Phillip. 



328 P. J. CHIMONIDES 

DESCRIPTION. In the erect part of the colony, branching is based on tri-, tetra- and pentafurcation. 
The colony is constructed of three types of stolons, here termed 'a', 'b' and 'c' (see below). Often, 
the region of branching, of a maternal stolon, is prominently thickened. Pentafurcation appears to 
be associated with external influence such as injury or the presence of an epibiont. Tetrafurcation is 
associated with astogenetically early regions, forming the base and main-axis regions of the colony. 
Main-axis regions are composed of type 'a' stolons. Trifurcation is associated with side branches 
which develop from main-axis regions. It is not possible to predict with certainty, the direction in 
which stolons will be produced at pentafurcation. In both the tetra- and trifurcate conditions 
however, one resultant component is produced in linear succession, and two others are produced 
laterally, one on each side. These are lateral side branches, composed of type 'b' stolons and are 
produced at an angle of about 60 deg. to the central axis. In the tetrafurcate condition, the fourth 
component, comprising type 'c' stolons, is produced posteriorly to its maternal stolon in the central 
axis, also at an angle of about 60 deg. This is a posterior side branch. Development along the side 
branches is limited and ends with the production of pinnately arranged, tapering kenozooids. The 
component kenozooids of such pinnate groupings are arranged as: three in linear succession, with 
an opposed lateral pair at both inter-kenozooidal junctions. In the lateral side branches, the 
pinnate kenozooids are usually produced after a 'linear' succession of three stolons; in the posterior 
side branch, after only one. The orientation of autozooids about the stolon along a main-axis 
sequence remains the same; this same orientation is preserved in the posterior side branch. In the 
lateral side branches, the autozooid group orientation is also preserved from stolon to stolon, but 
the autozooids are usually re-orientated to face distally along the main-axis; also in these branches, 
only one stolon, of a possible three, is usually produced at each branching point. On one side of the 
main-axis, viewed anteriorly, this is in the extreme right position; on the other side of the main-axis, 
this is in the extreme left. In each case, the other two positions are replaced by a pinnate terminal 
kenozooid group. The stolons along a lateral side branch are thus deflected anteriorly at each 
junction, in relation to the main-axis stolons. The branches therefore form inward facing arches 
across the anterior surface of the main-axis stolons. The resulting form is a long 'cylindrical' plume, 
reminiscent of snake vertebrae with ribs. The colony is composed of a number of these plumes, 
arising from various positions. Autozooid groups occur towards the distal ends of stolons. In the 
main-axis stolons, there is a further, distal, autozooid-free portion to each stolon, usually corre- 
sponding in length to the width of a daughter stolon. In the side branches, the autozooids fre- 
quently overlap the subsequent branching point. All stolons may be curved anteriorly. Autozooids 
are outer-wall thickened, but the thickening differential is usually low. Autozooid group profile 
tends to be level. The arrangement of autozooids in groups along main-axis stolons is, to some 
extent, predictable. A proximal-most autozooid may be evident in a group, and is associated with 
one side of the stolon. This autozooid loses its prominence over the next few stolons, the proximal 
autozoids of the groups appearing equally paired. Eventually, a proximal-most autozooid 
becomes prominent once more, but this time is associated with the opposite side of its stolon. Such 
a sequence is estimated to occur over 5 stolon units. The original condition is regained after a 
sequence of 10 stolon units. Side branches, where produced, have autozooid groups each with a 
prominent proximal-most autozooid associated with the side of its stolon nearest the main-axis 
stolons. This arrangement is preserved in subsequent autozooid groups along a side branch, unless 
a main-axis sequence is produced. Rhizoids may develop, one per stolon, from the proximal most 
end of, usually, main-axis stolons. Each rhizoid is produced at about 30 deg. to the orientation of 
the autozooids on the same stolon. 

SI. (a) 2-44 Z/S.(a)50% 

Zn. (a) (14-25 (appearing as 7-12 'pairs') 

SI. (b)l-60 Z/S.(b)80% 

Zn. (b) 18-28 (appearing as 9-14 'pairs') 

SI. (c)M3 Z/S.(c)80% 

Zn. (c) 18-28 (appearing as 9- 14 'pairs'). 



AMATHIA 329 

Sd. 0-35 (all stolons) 
Zh. 0-35 (all stolons) 
Zw. 0-1 3 (all stolons) 

REMARKS. The cuticle in some specimens is seen to bear numerous cyst-like bodies, whose structure 
and function have yet to be determined. These cysts appear to be associated with the distal, 
astogenetically later (most recently budded) parts of the colony. 

The branching pattern and resulting shapes in parts of the colony are quite distinct. The overall 
result is that autozooids face into a relatively sheltered colony-bounded space. This arrangement 
may have some protective advantage (see page 341). 

D'Hondt (1983) places A. verticillata Waters MS and A. delicatissima Busk MS in synonymy 
with A. wilsoni. Only the latter assertion is completely correct. The only apparent record of A. 
verticillata MS is of slide MM 7137, bearing the legend 'so named by Kirkpatrick . . . KP. after- 
wards called it Amathia wilsoni K'. A. verticillata is, thus, merely Kirkpatrick's MS name for what 
he subsequently described as A. wilsoni. The slide was part of Water's collection, from which the 
confusion probably arises. There appears to be no other record of A. verticillata Waters MS. 

In the original description by Kirkpatrick (1888), a BMNH specimen from Port Jackson is 
apparently indicated. No such specimen has been found. The entry in the account is somewhat 
anomalous, in that the account deals with 'Polyzoa from Port Phillip'. It seems likely that 
Kirkpatrick was referring to an additional specimen, then held in the collections at the BMNH, but 
whose whereabouts cannot now be determined, simply of the same identity as that which he 
described. At the beginning of the account, Kirkpartrick stated that he was describing new species 
from a collection made by J. B. Wilson from Port Phillip, subsequently sent to the BMNH. 
Specimen BMNH 1888.5.17.7 matches this description in being part of such a collection, and is 
indicated as type material in catalogue and registration records, in Kirkpatrick's own hand. This 
specimen is clearly syntype material. 

DISTRIBUTION. The species is known from the south-eastern region of Australia, ranging from 
Holdfast Bay near Adelaide to Port Jackson near Sydney and Hobart, Tasmania. The record from 
Flinders Island is not clear; it could refer to the island off Tasmania or that in the Great Australian 
Bight. 

Amathia pinnata Kirkpatrick, 1888 
(Figs3B, 10A,B) 

Amathia pinnata Kirkpatrick, 1888: 19, pi. 2 (figs 5, 5a). 

Amathia pinnata: MacGillivray, 1895: 136, pi. C (figs 1, la). 

Part Amathia pinnata: d'Hondt, 1979: 16. 

Part Amathia inarmata: d'Hondt, 1983: 67, fig. 36 (G). 

Not Amathia inarmata: d'Hondt, 1983: 67, pi. 2 (fig. \)( = A. biseriata). 

MATERIAL EXAMINED 

Lectotype (selected here): BMNH; 1888.5.17.8 A, Port Phillip, J. B. Wilson Collection. 
Paralectotypes: BMNH; 1888.5.17.8 B, C, Port Phillip, J. B. Wilson Collection. 

OTHER MATERIAL 

BMNH; 1 847.6.23. 14, Tasmania. 1 884. 1 1 . 14.5-1 2 B, Port Phillip. 1 886.6.8. 1 , Griffths Point, Port Jackson. 
1963.2.12.363, George Town, ?Tasmania? 1985.3.28.1, Port Phillip Heads, 15 m. 1985.3.30.2, Algoa Bay, S. 
Africa. 

MM; 7109/2W, Port Phillip. 

DESCRIPTION. In the erect part of the colony, branching is regular and almost always trifurcate. 
This results in a typically compound pinnate arrangement. At any branching point, one daughter 
stolon is produced in rectilinear succession, and two others are produced laterally opposing, at an 
angle of about 60 deg. to the centre. Stolons tend to be straight. Autozooids are distally located, 
occupying the greater part of stolons. Autozooid groups develop as far as the subsequent branch- 
ing point, but do not overlie it. Autozooid group profile tends to be level, and autozooids are 
outer-wall thickened, although there is a tendency for both differential and overall thickening, not 



330 P. J. CHIMONIDES 

to be great. Autozooid group orientation from stolon to stolon is generally well preserved. The 
arrangement of autozooids on stolons lying in rectilinear succession is to some extent predictable. 
Viewed anteriorly, a proximal-most autozooid may be prominent in an autozooid group, and is 
associated with one side of the stolon. Over successive stolon units, each autozooid group shows 
rearrangement so that this autozooid loses prominence. The proximal autozooids thus appear 
equally paired, until a proximal-most autozooid becomes prominent once more, this time 
associated with the opposite side of the stolon. Such a series appears to occur over a sequence of 4 
stolon units. The original condition is recovered on the seventh or eighth stolon unit. In lateral 
daughter stolons, a single proximal-most autozooid is prominent in the autozooid group, this 
being associated with the side of the stolon nearest the rectilinear series, i.e. for a right-branched 
stolon, the proximal-most autozooid is nearest the left side of its stolon, and vice versa. 
Autozooid groups on stolons subsequently produced from a lateral daughter stolon, display the 
same organisation along resultant rectilinear series and lateral components. Terminal lanceolate 
processes may occasionally be produced; each one is made up of a tapering series of three 
kenozooidal sub-units. These terminal lanceolate processes are usually produced simultaneously 
as a group of three, each process replacing a normal stolon. Rhizoids may be produced, 
approximately mid-way along the proximal autozooid-free part of the stolon. These arise singly 
or as a pair, one on either side of the stolon, at about 30 deg. to the autozooid orientation. 
Colonies may be large and arborescent. 

Zh. 0-45 Z/S.80% 

Zw. 0-13 Zn. 18-49 (appearing as 9-24 'pairs') 

SI. 1-50-3.25 Tpl. 1-50 

Sd. 0-35 (just proximal to the autozooid group) 

REMARKS. Measurements of this species given by Kirkpatrick (1888) appear to originate from the 
same material as he figured. There is, however, some discrepancy between the figures and the 
description, as it is possible to infer 18 autozooid 'pairs' from his figure, whereas he described 
the range as being from '12-16'. 

Kirkpatrick's figure corresponds to a specimen which is obviously a fragment from a larger 
colony. However, this specimen and another which greatly resembles it, are obviously not from 
other material stored in the same container and bearing the same registration number. All these 
specimens are Kirkpatrick's A. pinnata, as is borne out by catalogue and registration records in 
Kirpatrick's own hand. There is a suggestion, in the stolon shape and rhizoid production site, that 
the registration may harbour two species. A. pinnata sensu stricto, is taken as the morph which 
corresponds with Kirkpatrick's figure; the registration of these components receiving the suffixes A 
and B (the remaining component the suffix C). Component A is the figured specimen, and is here 
designated the lectotype, the remaining portions, B and C, being paralectotypes. Provisionally, all 
three components are accepted as being A. pinnata. 

This species is one of a number that were considered by d'Hondt (1979, 1983) to be synonymous 
with one another, the grouping also including: A. biseriata; A. tricornis (part); A. brongniartii; A. 
cygnea MS; A. 'polycistica' MS; A. desmarestii MS (see page 331). D'Hondt (1979) indicated A. 
pinnata Kirkpatrick 1888, as the senior synonym of this group. D'Hondt (1983) then indicates that 
A. inarmata MacGillivray 1887, is the senior synonym of the same compositional group, thus 
subordinating A. pinnata as a junior subjective synonym. A. pinnata is in fact not synonymous with 
any of the species in this grouping, being a separate and distinct species (see pages 332, 333). 

DISTRIBUTION. The species is known from the south-eastern region of Australia, ranging from Port 
Jackson, to Port Phillip Heads and Tasmania, also being recorded from Algoa Bay, South Africa. 

A mat hia biseriata Krauss, 1837 
(Figs4B, 11C,D) 

Amathia biseriata Krauss, 1837: 23, fig. 1 (a, b, c). 

Not Amathia biseriata: Busk, 1852: 385. 

? Amathella biserialis Gray 1858: 320 (? errorum pro Amathia biseriata Krauss, 1837). 



AMATHIA 331 

Amathella uniserialis Gray, 1858: 320. 

Amathia inarmata MacGillivray, 1887: 183. 

Amathia biseriata: Kirkpatrick, 1888: 17. 

Amathia inarmata: MacGillivray 1889: 309, pi. 183 (fig. 4). 

Amathia biseriata: MacGillivray, 1895: 137, pi. B (fig. 4). 

part Amathia pinnata: d'Hondt, 1979: 16. 

part Amathia inarmata: d'Hondt, 1983: 67, fig. 36 (G). 

MATERIAL EXAMINED 
Neotype (selected here): BMNH; 1887.12.10.90, Port Phillip, J. B. Wilson collection. 

OTHER MATERIAL 

BMNH; 1899.7.1.4317,4318, 1963.2.12.357, Australia. 1899.7.1.4319, New Zealand. 1965.8.12.19, Jervis 
Bay, Huskisson, N.S.W. 1985.3.14.1, no locality. 

NMV; H492 (1-9), Port Phillip Heads. 

RM; 1808, Port Natal, Africa. 

DESCRIPTION. In the erect part of the colony, branching is always bifurcate. Viewed anteriorly, at 
each branching point, one daughter stolon tends to lie approximately in line with the maternal 
stolon, although deviations of up to 20 deg. may occur. The other daughter stolon is usually 
produced laterally, at an approximate angle of 45 deg. to the main axis of the maternal stolon. The 
side on which a lateral branch appears, alternates at each successive bifurcation. Stolons have 
a slight constriction near the proximal end and are usually curved anteriorly, the curvature 
becoming progressively more acute distally, to bend around the distal end of the autozooid groups. 
The stolon tends to remain in contact with the distal side of the autozooids. Where this is not so, 
this portion of the stolon remains autozooid-free. Daughter stolons produced in the 'linear' 
position, arise from the distal end of the maternal stolon; lateral daughter stolons are produced 
from the most sharply curved region of the maternal stolon. The distal region of the maternal 
stolon may show some axial subdivision to bear the daughter stolons, more so when it is not in 
contact with the distal face of the autozooids. Autozooid groups on maternal stolons are rarely 
developed distal to the origin of the lateral daughter stolon, or where the stolon shows division. 
Autozooids are outer-wall thickened, the thickening sometimes being accentuated at the rims. 
Autozooid group profile appears level, sometimes slightly concave centrally, or diminishing 
slightly proximodistally along the stolon. A single proximal-most autozooid is evident in each 
group, slightly broader than the rest, usually placed just off the mid-line, and thus associated with 
one or other side of the stolon. No pattern has been discerned in the location of this autozooid from 
stolon to stolon. Sometimes both daughter stolons show the same autozooid arrangement as on 
their maternal stolon; sometimes the opposite; sometimes combinations of the two. Autozooid 
orientation is generally well preserved from stolon to stolon. Rhizoids may be produced, one per 
stolon, arising from a position level with, or just proximal to, the proximal-most autozooid of the 
autozooid group. The orientation of the rhizoids is between 90-135 deg. to the autozooid group, 
occurring on the same side of the stolon as the direction in which that stolon was budded. Stolons 
and their autozooid groups become shorter nearer the distal (astogenetically later i.e. most recently 
budded) regions of the colony. 

SI. 2-05 Z/S.75% 

Sd. 0-32 Zn. 5-25 (appearing as 4-1 2 'pairs') 

Zh. 0-35 

Zw. 0-11 

REMARKS. The autozooid pattern from stolon to stolon remains elusive in this species, primarily 
due to the difficulties of observation over the number of stolon sequences necessary. 

This species is one of a number that were considered by d'Hondt to be synonymous with one 
another. D'Hondt (1979) indicated that A. pinnata Kirkpatrick 1888, was the senior synonym of 
this group, inclusive of A. biseriata, and then (1983) indicated that A. inarmata MacGillivray 1887 
was the senior synonym of the same compositional group. Both assertions are erroneous. The 
inclusion of the name A. desmarestii in this group is of little consequence as Lesueur never 



332 P. J. CHIMONIDES 

published his work. The name itself is not valid in being published only in synonymy (I.C.Z.N. 
article 1 le). Of the other species in this group: A. tricornis is a separate and distinct species (see 
page 321), with a more complex colony composition than the rest; A. cygnea Busk MS, and 
A. polycistica (sic) Busk MS, are here considered to be synonymous with one another as 
A. brongniartii (see page 333); A. pinnata is also a separate and distinct species, in which 
trifurcate branching predominates. There are no indications, in any of MacGillivray's accounts 
of A. biseriata, of the trifurcate branching pattern shown by d'Hondt (1983), who reproduced 
MacGillivray's (1895) figure of A. pinnata. In fact, MacGillivray and Krauss both stated that the 
branching pattern in A. biseriata is dichotomous. Supplementary features which may be used to 
distinguish between A. biseriata and A. pinnata are: the site of rhizoid production; the occurrence of 
terminal kenozooids in the latter species. Were A. biseriata and A. pinnata synonymous, then 
A. biseriata would be the senior synonym (cf. d'Hondt 1979). A. biseriata however, is synonymous 
With A. inarmata, but again, it is A. biseriata which is the senior synonym. In this case, Krauss' 
publication predates that of MacGillivray by 50 years. Furthermore, MacGillivray (1895) 
accepted his species to have been the same as that of Krauss. MacGillivray's syntypes are held in 
the NMV (H492 1-9), and all 9 specimens are A. biseriata. 

MacGillivray (1895) appears to have been under the misconception that Krauss' material of 
A. biseriata originated from south Africa, when in fact it was from New Holland i.e. western 
Australia. It is probable that MacGillivray (1895) was actually referring to material received from 
'Pergens' (MacGillivray, 1889). 

Krauss' (1837) description and figures are here considered to be more than adequate to dis- 
tinguish his species from any other; his only mistake was to make the assumption that the rhizoids 
produced the stolons and their autozooids. Krauss' material does not appear to have survived. In 
view of the subsequent confusion, selection of a neotype is necessary. BMNH 1887.12.10.90 
(although from Port Phillip) is selected here. 

DISTRIBUTION. The species is known from south Africa, southern Australia and New Zealand. 

Amathia brongniartii Kirkpatrick, 1888 
(Figs4A, 11A,B) 

Amathia brongniartii Kirkpatrick, 1888: 18, pi. 2 (figs 3, 3a). 

Amathia brogniartii (sic): lapsus calami MacGillivray, 1 895: 1 36, pi. B (figs 3, 3a). 

Part Amathia pinnata: d'Hondt, 1979: 16. 

Part Amathia inarmata: d'Hondt, 1983: 67. 

Not Part Amathia inarmata: d'Hondt, 1983: 67, fig. 36G, ( = A. pinnata). 

MATERIAL EXAMINED 
Neotype (selected here): BMNH; 1888.5.17.6, Port Phillip, J. B. Wilson Collection. 

OTHER MATERIAL 

BMNH; 1838.2.26.13, 1847.6.23.4, 1899.7.1.4379, 4381, Tasmania. 1887.4.27.19, Port Jackson, N.S.W. 
1887.12.10.98A (part), Port Phillip, Viet. 1899.7.1.4419, 6601, Swan Is. ?Bass Strait? 1927.9.26.21, Swan Is., 
Banks Strait. 1984.12.4.1, Portsea Pier, Victoria, 2m. 1985.3.16.1, Victoria. 1985.3. 16.1.2a,b, Flinders Is. 
TTasmania? 

NMV; 65397, Port Phillip Heads, Viet. 

MM; 7074, Lane Cove, Port Jackson, N.S.W. 

DESCRIPTION. In the erect part of the colony, branching is always bifurcate. At each branching 
point, one daughter stolon tends to continue in line with the maternal stolon, often giving rise to 
rectilinear series, although deviations by up to 15 deg. may occur. The other daughter stolon of 
each birfurcation arises laterally, often anterolaterally, appearing on alternate sides along a series, 
at an angle of between 20-50 deg. to the axis of the maternal stolon. Stolons are usually straight and 
tend to have a slight constriction near the proximal end. The distal end of a stolon does not usu- 
ally show any axial subdivision or widening to bear daughter stolons; more often, the maternal 
stolon shows some abbreviation into a wedge shape to accommodate these. The autozooid groups 
develop as far as, and often overlie the subsequent branching point. Autozooids are markedly 
inner-wall thickened, with a thin walled exterior. The autozooids are usually, large, prismatic, and 



AMATHIA 333 

pentagonal in section. Viewed anteriorly, the walls between the autozooids, being much thicker, 
give a characteristic zig-zag backbone appearance to the autozooid groups. Autozooid group 
profile ranges from gently arched upwards to level with the autozooids shorter at each end of the 
group. A single proximal-most autozooid is evident in each autozooid group, slightly broader than 
the rest, usually just off the mid line and thus associated with one or other side of the stolon. Viewed 
anteriorly, this autozooid is always on the side nearest the sister stolon. Autozooid orientation 
from stolon to stolon is generally well preserved. Rhizoids may be produced, one per stolon, from a 
position level with or just proximal to the proximal-most autozooid of a group. These arise at 
about 1 10-160 deg. to the autozooid orientation on the same stolon, and on the same side of the 
stolon as the direction in which the stolon was budded. Stolons and their autozooid groups appear 
to be shorter nearer the distal (astogenetically newer) regions of the colony. 

SI. 1 -75^-00 Z/S.80% 

Sd. 0-25 Zn. 10-39 (appearing as 5-18 'pairs') 

Zh. 0-48 

Zw. 0-15 

REMARKS. This species was considered by d'Hondt to be a junior synonym of: (1979) A. pinnata; 
then (1983) of A. inarmata. A. brongniartii differs from A. pinnata in many features, such as: the 
autozooidal thickening; the sites of rhizoid production; the basic branching pattern. A. inarmata is 
itself a junior synonym of A. biseriata (see page 332). Kirkpatrick (1888) and MacGillivray (1895) 
indicated differences between A. biseriata and A. brongniartii in their accounts. The two species 
may be distinguished quite readily by: the shape of the stolons; to some extent, the site of rhizoid 
production; the autozooidal thickening, this last being the most prominent difference. 

A. brongniartii appears to display a large variation in stolon length and attendant number of 
autozooids borne. Such variation may be seen within single colonies. However, colonies may often 
show good uniformity in stolon lengths, whether long, short or intermediate. Busk, in his unpub- 
lished notes and figures stored at the BMNH, considered the possibility that the extremes of the size 
range might be discrete. He appears to have called colonies with short stolons and lower autozooid 
number A. cygnea (up to 20 autozooids, equivalent to 8-12 'pairs'), with more diminutive versions 
as A. cygnea var. nana. Colonies with higher numbers of autozooids (24-36 units, equivalent to 
12-18 'pairs') and longer stolons, he called A.polycystica. In Busk's material, the specimens which 
might be A. polycystica tend to be dark coloured, but other than this there seems to be nothing 
which distinguishes them taxonomically. (Busk's notes make no recognition of the A. brongniartii 
in the unpublished plates of Lesueur). 

Kirkpatrick (1888), in his account of A. brongniartii, erroneously credited the species to 
Desmarest and Lesueur, citing Lesueur's figures and Pergens' (1887) collations as his reference for 
the identity and name. As Desmarest and Lesueur never published their work, the Pergens men- 
tions the name only in synonymy, Kirkpatrick is the authority for the species (I.C.Z.N. article 1 le). 
As Kirkpartick did not consider himself the author of the species, he did not choose any type 
specimens. Kirkpatrick's figures, like those of Lesueur, are of insufficient quality to be utilised as 
reliable references. Kirkpatrick's figures hardly show any detail at all, whilst Lesueur's figures 
(pi. 13 fig. 5) show curved stolons and tube-like autozooid anteriors, both characteristics of 
A. biseriata. At the same time however, Lesueur shows approximately 20 'pairs' of autozooids, a 
number high enough to be associated with A. brongniartii. Understandably, confusion has arisen 
and so it would seem appropriate that a neotype be designated. BMNH 1888.5.17.6 is, therefore, 
selected as neotype.- This specimen is the only one labelled as A. brongniartii by Kirkpatrick in the 
collection made by J. B. Wilson from Port Phillip. This collection is the subject of Kirkpatrick's 
publication of 1888. 

Specimen NMV 65397 is believed to have been before MacGillivray at the time of his writing his 
1895 account of Amathia species (in litt. NMV. 30th May 1983.). The specimen is undoubtedly A. 
brongniartii Kirkpatrick, thus MacGillivray's name 'A. brogniartii' is simply a misspelling, as his 
synonymy indicates. 

DISTRIBUTION. The species is known from south-eastern Australia, ranging from Tasmania to Port 
Jackson. 



334 P. J. CHIMONIDES 

Amathia alternata Lamouroux, 1816 
(Figs4C, 13A,B) 

Amathia alternata Lamouroux, 1816: 160. 

Amathia alternata: Lamouroux, 1821: 10, pi. 65 (figs 18, 19). 

Amathia alternata: Lamouroux, 1824: 44. 

Not Amathia alternata: Osburn, 1932: 444, pi. 1 (fig. 4). 

Part Amathia convoluta: Mature, 1957: 22, fig. 11. 

Not part Amathia convoluta: Mature, 1957: 22, fig. 10. 

Amathia alternata: Winston, 1982: 108, fig. 8. 

MATERIAL EXAMINED 
Neotype (selected here): USNM; 6307 (part), Albatross Stn. 2619, off Cape Fear, North Carolina. 

OTHER MATERIAL 

BMNH; 1964.7.10.1 A,B, New River Inlet, North Carolina. 
1964.7.10.2, Alligator Harbour, North Carolina. 

DESCRIPTION. In the erect part of the colony, branching is always bifurcate. At each branching 
point, one daughter stolon tends to continue in line with the maternal stolon, forming a linear 
series. The other daughter stolon is produced at approximately 45 deg. to the maternal stolon axis 
at that location, at between 45-90 deg. to the orientation of the distal autozooids there. Branching 
may appear equally dichotomous at times. Daughter stolons are produced from the posterior side 
of the maternal stolon, this showing some abbreviation into a wedge shape to accommodate them. 
Stolons are: narrowed proximally, additionally having a proximal constriction; often curved 
posteriorly, also undergoing a slight twist along their length. The linearly disposed stolons may 
thus appear as an undulating progression. Any twist in these is normally reflected in the autozooids 
borne, changing the orientation between proximal and distal autozooids in a group by up to 90 deg. 
At times, this may give the impression that autozooid groups are simply arranged obliquely on the 
stolons. Autozooid groups may, however, be arranged along the stolonal axis without any evi- 
dence of twist at all in either component. Where the twist, proximodistally along a stolon, is 
clockwise, the left daughter stolon is produced in the 'lateral' position, and with anticlockwise 
twist, the right. The direction of twist is generally well preserved from stolon to stolon (although 
both directions may be found in the same colony). Lateral daughter stolons are thus produced 
from the same side of stolons along any linear sequence. The spatial orientation of these lateral 
branches is determined by the maternal axis and autozooid orientation at that point. Autozooid 
orientation changes by 100-1 80 deg. from stolon to stolon. This is taken from the distal autozooids 
on the maternal stolon to the proximal autozooids on each of the two daughters. Autozooids 
frequently overlie the subsequent branching point and are outer- wall thickened. Autozooids are 
inclined at about 60 deg. to the stolonal axis, and group profile is level or gently convex. A 
proximal-most autozooid is frequently evident in each group. Viewed anteriorly, this is associated 
with the side of the stolon in which the direction of twist occurs e.g. the right side, with clockwise 
twist proximodistally. Rhizoids may be produced at any point on a stolon from the constriction to 
beneath the proximal autozooids. A polyrhizoid condition may result, where any number of 
rhizoids, up to a maximum of 5, possibly more, may be produced from a single stolon, at any 
orientation. Two orientations appear more frequently occupied by rhizoids: within 10 deg. of the 
proximal autozooid orientation on the same stolon; approximately 180 deg. to the proximal 
autozooid orientation. 

Zh. 0-42 Z/S.85% 

Zw. 0-10 Zn. 24-57 (appearing as 12-28 'pairs') 

SI. 1-45-3-87 

Sd. 0-29-0-35 (just proximal to the autozooid group) 

REMARKS. In the non-erect part of the colony, secondary thickening of stolons may occur. This has 
the appearance of a sleeve developing along existing stolons. 

A. alternata can display a consistent, if only slight, spiral nature, and the species has been 
confused with A. convoluta sensu Lamouroux, and possibly also with A. semiconvoluta 



AMATHIA 335 

Lamouroux. As no type material exists, it may also be possible to confuse A. alternata with other 
spiral-autozooid group species, for example A. tortuosa Tennison Woods or A. connexa Busk. 

Mature (1957) described and gave representative figures, under the name of A. convoluta (sensu 
Lamouroux), of material collected at Fort Macon on Bogue Banks, USA. This material was noted 
to have 'straight autozooid groups, alternately placed from one internode to the next', and to be the 
same as a specimen in the USNM labelled A. spiralis, from Albatross Stn. 2619, off Cape Fear, 
North Carolina. The latter, USNM 6307 (part), has been examined here. Portions of this specimen 
have autozooids arranged and placed in the way Mature describes and illustrates for the Fort 
Macon material. This specimen, Maturo's account, and the description given here, conform with 
Lamouroux's brief description of A. alternata in 1816, his subsequent account of 1821, in which he 
presents figures (pi. 65, figs 18, 19), and his final account in 1824. 

Lamouroux's (1821) figures lack important information, and, in not being published at the same 
time as the original description, are clearly not eligible for any type status. Lamouroux's collection 
was destroyed during the Second World War (d'Hondt in litt. 27.10.1982), but material from his 
collection obtained via Busk, and labelled A. alternata is stored at the BMNH as 1897.7.1.6606. 
This material was originally stored dried and pressed, but has subsequently been rehydrated, and is 
at present stored in alcohol. As recorded on a label with the material, examination by Dr F. Mature 
before rehydration revealed only two bryozoan species, these being other than A. alternata, 
possibly A. brasiliensis Busk and Zoobotryon verticillatum. Since rehydration, the absence of A. 
alternata is here confirmed, and the identity of the two other species established as A. wilsoniand A. 
semiconvoluta. 

The packet originally enclosing the specimens bears the names 'A. alternata' and 'Amerique' in 
what is taken to be Lamouroux's handwriting. The locality mentioned does not disagree with that 
of Lamouroux's accounts (1816, 1821, 1 824) of A . alternata, (the most specific locality given being 
the Sea of Antilles in 1824). The two species found enclosed in the packet, however, are not 
expected from this region; all other records of A. wilsoniare from southern Australia. Similarly, all 
other records for A. semiconvoluta are from the Mediterranean, the species possibly extending as 
far along the west African coast as Nigeria. It is unlikely that A. wilsoni would have been confused 
with A. alternata by Lamouroux, as the species has many distinguishing features and lacks alter- 
nate autozooid group placings. It may be possible to confuse dried A. semiconvoluta with A. 
alternata, but this is considered unlikely of Lamouroux, as he is the author of both species. 
Lamouroux introduced A . alternata in 1 8 1 6, redescribed the species in 1 82 1 and again in 1 824 when 
he introduced A. semiconvoluta, the descriptions for the two species appearing on the same page. It 
is to be assumed that the author was capable of recognising and distinguishing his own species. The 
fate of any A. alternata that may have been present in BMNH 1897.7. 1 .6606 is open to speculation. 
The circumstances of Busk's acquisition of specimens enveloped in paper bearing Lamouroux's 
writing are unknown. 

In the interests of nomenclatural stability, a neotype is required. Specimens considered eligible 
are: those in the BMNH under 1964 registrations, all from North Carolina, donated and identified 
by Dr F. Mature as A. alternata; specimen USNM 6307 (part); Maturo's Fort Macon material. 
The whereabouts of the Fort Macon material (Mature 1957) is not known. The neotype selected, 
therefore, is specimen USNM 6307 (part) from Cape Fear, off North Carolina, at 1 5 fthms. 
(27.43 m) this being the earliest recorded specimen surviving. The polyrhizoid condition is not 
readily apparent in this specimen, but it does show the possible variation in the arrangement of 
autozooid groups. 

The difference between A. alternata and A. semiconvoluta are as follows: the degree of 'spirality' 
that may occur is much greater in A. semiconvoluta (180-270 deg., cf. A. alternata 0-90 deg.); the 
orientations of the distal end of one autozooid group and the proximal end of the next are within 10 
deg. of each other in A. semiconvoluta, but a distinctive 100-180 deg. in A. alternata; a lower linear 
autozooid to stolon ratio of 50% for A. semiconvoluta, compared to about 85% in A. alternata. 
Although rhizoids may appear at similar orientations in both species, only A. alternata shows 
polyrhizoidy with rhizoids in proximity to the autozooids (and, additionally, a slightly wider 
bifurcation angle between daughter stolons). In A. semiconvoluta the rhizoids appear at the 



336 P. J. CHIMONIDES 

proximal-most end of the stolons, at about the same orientation at which the preceeding autozooid 
group terminates, or displaced by 1 80 deg., or when two rhizoids are present on the same stolon, at 
both orientations. A. semiconvoluta is understood from the following specimens: BMNH; 
1885.12.5.12,13, Marseilles. 1888.11.9.4, Naples. 1899.5.1.290, 1912.12.21.687, Adriatic. 
1 899.7. 1.6606pt., locality? 

The other species mentioned above i.e. A. convoluta, A. tortuosa and A. connexa, are also 
distinguishable from A. alternata by their degree of spirality. This is significantly greater than A. 
alternata in all cases. Problems might arise, however, in distinguishing these three species from 
each other, and establishing their validity. 

A. convoluta is understood from BMNH 1899.7.1.6607. This specimen is from Lamouroux's 
collection, obtained via Busk, and is labelled 'Amathia convoluta, Australasia' in what is 
accepted to be Lamouroux's handwriting. There is nothing to contradict its identity from any of 
Lamouroux's descriptions. The specimen also conforms with MacGillivray's (1 895) account of the 
species, corroborated by his opinion on Busk 1884 (pi. 6. fig. 2, there misidentified as A. spiralis). 
However, there is nothing to suggest that MacGillivray ever saw BMNH 1899.7.1.6607 at any 
time. This specimen is noted as 'type' in the catalogue of the BMNH, though no formal declaration 
of its purported status has ever been made. It is possible that the specimen was formative of 
Lamouroux's opinions of the species and thus a 'type' but there can be no certain evidence for or 
against this notion. However, the specimen appears to be the only extant material which bears an 
unchallengeable identification, attributable to the original author. D'Hondt (1983) indicated that 
Lamarck's name for the species (Amathia crispa), as the senior synonym, should instead be used. 

DISTRIBUTION. The species is recorded off North Carolina, USA, and, from Lamouroux's (1824) 
record, from the Caribbean. 

Amathia pruvoti Calvet, 1911 
(Fig. 13D) 

Amathia pruvoti Calvet, 191 1: 59, fig. 2. 
Amathia pruvoti: Bobin & Prenant, 1956: 287, fig. 128. 
Amathia pruvoti: d'Hondt, 1983: 67, fig. 35F. 
Amathia pruvoti: Hayward, 1985: 136, figs 46A, B. 

MATERIAL EXAMINED 
Type: LBIMM; Bry 8205, Calvet collection: no locality. 

OTHER MATERIAL 

BMNH; 1882.7.7. 1-2, Trieste. 1882.7.7.-, Mediterranean. 

1885.12.5. 14, Montpellier. 1889.7.27.48, 1890.7.22.8 part, Studland Bay, Dorset. 1975.7. 1.1 5, Emborios Bay, 
Chios, 90 ft. 1984.2.26.102, Dhiaporia Rock, Chios, 100ft. 

DESCRIPTION. In the erect part of the colony, branching is always bifurcate. Daughter stolons 
appear to diverge equally, lying at approximately 60 deg. to each other, thus giving the impression 
of equal dichotomy. In fact, at each branching point, one stolon tends to be budded in a linear 
position and is subsequently deflected, whilst the other is produced laterally. The linearly dis- 
posed daughter stolon may be wider than its sister, with little deflection, at times giving a strong 
impression of rectilinear progression. In all stolons, there is a slight constriction near the proximal 
end. The distal end does not show any axial subdivision or widening to bear daughter stolons; 
however, it often shows some abbreviation into a wedge shape to accommodate these. Autozooid 
groups sometimes overlie the subsequent branching point. However, it is more usual for the 
autozooid group to only develop as far as the branching point, or, alternatively, 'stop short' and be 
followed by an autozooid-free portion of stolon, about the width of an autozooid in length. Stolons 
tend to be straight proximally although often slightly curved posteriorly and undergoing an axial 
twist in the region of the autozooids. The twist in the stolon is usually reflected in the autozooids 
borne, changing the plane of their orientation, between the proximal-most and distal-most auto- 
zooids, by approximately 90 deg. The plane in which the subsequent bifurcation occurs is also 



AMATHIA 337 

affected to the same degree. The twists are normally predictable. Viewed anteriorly, in the left- 
branched daughter stolon, the twist is usually clockwise in a proximodistal direction, and anticlock- 
wise in a right-branched daughter. These twists generally occur irrespective of the twist which 
occurs in the maternal stolon. However, there can be variations to this. Occasionally, both 
daughters may twist in the same direction, this being opposite to that of their maternal stolon. 
Occasionally, the inverse to the normal condition occurs, where a left daughter twists anticlockwise 
and the corresponding right daughter twists clockwise. Autozooids are outer-wall thickened. 
Autozooid group profile, where discernible, is level proximally, diminishing distally, resulting 
from decreasing height and increasing distal inclination of the autozooids. A single proximal-most 
autozooid is evident in each autozooid group, usually off centre to the axis of the stolon. Viewed 
anteriorly, this autozooid is always associated with the same side of the stolon, as the direction 
in which the autozooid group twists, e.g. the right side, with clockwise twist proximodistally. 
Autozooid orientation from stolon to stolon, changes by 1 80 deg. between the distal autozooids of 
the maternal stolon and the proximal autozooids of each of the two daughter stolons. No rhizoids 
are known, and the erect part of the colony appears as a diffuse cotton- wool like mass. Sometimes, 
erect components of the colony come into contact with the substratum, and their characteristic 
stolonal shape is lost. These components do not bear autozooids; as stolonal kenozopids (see 
page 309), they become elongated and twisted, occasionally branching and producing clumps of 
flattened lateral processes. Further erect components may be produced at any time and these may 
resume the normal erect growth pattern. 

SI. 2.40-3.75 Z/S.60% 

Sd. 0-13-0-15 Zn. 21-31 (appearing as 10-15 'pairs') 

Zh. 0-40 

Zw. 0-13 

REMARKS. Apart from the ancestrula, little is known of the non-erect portion of the colony. It is 
assumed that this would resemble the contact-modified erect stolons and their growth behaviour. 
No occurrence of two autozooid groups on the same stolon has been encountered in any of the 
material examined (cf. Calvet, 1911). Such an instance would be contrary to the present concept of 
the genus. 

There is a specimen at the LBIMM, bry 8205, originating from the Station Zoologique de Cette, 
Universite de Montpellier. This is latterly documented (e.g. LBIMM loan form 26th Oct 1983) as 
'the probable type of A. pruvoti: Calvet (Cette), with a handwritten label of the author carrying the 
name A. semiconvolutcC . The justification for regarding LBIMM bry 8205 as the type specimen of 
A. pruvoti Calvet, is not given. The specimen is, however, well preserved, and would serve as an 
excellent basis on which to recognise the species in future. It is proposed here that the specimen be 
accepted as the type specimen of the species. If no historical justification for its claimed status as a 
'type' is available (see below), it is here selected as neotype, obviating the confusion that has arisen 
between A. pruvoti and A. lendigera (sensu lato). 

It should be noted that there is some difference between Calvet's description (1911) and 
specimens subsequently recognised as A. pruvoti, including specimen LBIMM bry 8205. Calvet 
described stolons as lying in rectilinear series. This condition is not readily apparent in the majority 
of specimens, except in two specimens from Chios, BMNH 1 975.7. 1.15,1 984.2.26. 1 02, and in these 
there is also little evidence of the proximal stolonal constriction. It is not possible to be certain of 
what Calvet meant when he described the 'stature' of A. pruvoti as 'erect', then drawing a compara- 
tive difference between it and A. lendigera, when the colony budding patterns of the two species are 
in fact very similar. It is possible that the supposed distinction may reflect an opinion that A. 
lendigera has a higher proportion of the non-erect colony component, or that the erect part of 
A. lendigera tends to be spatially more condensed. In both species, there is some variation in the 
overall length of stolons. This variation appears less extensive in A. pruvoti. The most obvious 
difference between the two species, however, lies in the disposition of autozooids about the stolons. 
A degree of twist is usually present in A. pruvoti, and an autozooid-free distal portion of the stolon 
often occurs. 



338 P. J. CHIMONIDES 

Calvet also drew a comparison with A. semiconvoluta. The differences in the erect part of the 
colony between this species and A. pruvotiare that, in A. semiconvoluta: the curvature of the stolon 
beneath the autozooids is much shallower, if present at all; the autozooid height tends to be equal 
throughout the autozooid group; the autozooid group is more spiralled, undergoing twists of 
180-270 deg.; the orientations of the distal end of the autozooid group on the preceeding stolon, 
and the proximal ends of the next, on the succeeding stolons, occur within 10 deg. to each other; 
the direction of spiral tends to be preserved from maternal to daughter stolons, although both 
directions may be found in the same colony; branching is always bifurcate (as in A. pruvoti) but one 
daughter stolon is always linearly disposed, giving rise to definite rectilinear series, with the other 
daughter stolon produced anterolaterally at about 30 deg. to the stolon axis and distal autozooid 
orientation; when the autozooid twist, proximodistally, is clockwise, the right hand daughter 
stolon is in the rectilinear position, and with anticlockwise twist, the left hand daughter stolon 
acquires the rectilinear position; autozooid groups always overlie the subsequent branching 
point; rhizoids are produced from the proximal end of stolons. Further characteristics of A. 
semiconvoluta are as follows: rhizoids arise singly, either in the same orientation as the proximal- 
most autozooids, or at 180 deg. to this (see pages 335, 336, Figs 5A, 13C); when two rhizoids per 
stolon are produced, these arise as one from each orientation; the production of rhizoids would 
enable the colony to attain an arborescent form, but this has not been confirmed. 

There is, in addition, some similarity between A. pruvoti, A. distans Busk, A. distans var. 
aegyptana d'Hondt and A. brasiliensis, each of which is a distinct entity. The distinction between 
the species may be found in the following characteristics. In the last three, the autozooid groups are 
more spiral, usually describing a 360 deg. rotation about the stolon in A. brasiliensis and A. distans, 
slightly less (270-360 deg.) in A. distans var. aegyptana. Of this group, A. brasiliensis is the only one 
which produces rhizoids, these arising at the proximal end of stolons, orientated within 10 deg. to 
the proximal autozooids on the same stolon. A. distans var. aegyptana has the distinction of 
producing autozooid groups in which the direction of spirality remains preserved from maternal to 
daughter stolons i.e. all clockwise or all anticlockwise, whereas one of two other patterns prevail in 
A. pruvoti, A. distans and A. brasiliensis. Using the distal-most autozooids as the orientation 
reference, and viewing anteriorly: in A. pruvoti and A. brasiliensis, the left daughter stolons carry 
autozooids arranged clockwise in a proximodistal direction, and the right daughters, anticlock- 
wise; in A. distans, the left daughter stolons carry autozooids arranged anticlockwise, and the right 
daughters, clockwise. The distinctions are made with reference to type material: 

For ,4. distans: BMNH 1887.12.9.926, Bahia, 10-12 fthms. (18.29-36.58 m.). 
For A. brasiliensis: BMNH 1887.12.9.927, Bahia, 10-20 fthms. (18.29-36.58 m.). 
For A. distans var. aegyptana: BMNH 1926.9.6.25, Suez Canal. 
For A. pruvoti: LBIMM bry 8205, no locality. 

In conclusion, A. distans var. aegyptana should be considered as a species in its own right, and is 
here raised to specific rank as Amathia aegyptana. 

Harmer (1915) drew attention to the similarities between A. distans and other species, including 
A. pruvoti. However, his understanding of A. distans, particularly in the degree of spirality which 
may occur, is here considered insufficiently rigorous. Unfortunately, it is Harmer's understanding 
which is followed by Bobin and Prenant (1956) and d'Hondt (1983). 

DISTRIBUTION. The species is known mainly from the Mediterranean, with some material from 
Studland Bay in Dorset, England. 



Discussion 

It is readily apparent that there is a considerable degree of regularity and possible colony inte- 
gration within species of the genus Amathia. Some of this is reflected in the consolidation of a 
colony by rhizoids. These grow back, sometimes fusing with each other, and ultimately interact 



AMATHIA 339 

with the substratum to provide support. The various arborescent growth forms that result can only 
be maintained through continued sustenance of these rhizoids, and of any underlying stolons 
which will usually have lost their feeding autozooids. This implies nutrient transfer to them, and 
thus a potential ability for self repair. 

The most basic and obvious level of intergration, however, is the clustering of autozooids into 
groups on septa-bound kenozooidal stolons, to form intercommunicating functional units. These 
can show changes of characteristics with astogeny. Changes may be gradual, as in stolon lengths 
and autozooid numbers in A. biseriata; or discontinuous, as in the autozooid complement per 
stolon in parts of A. tricornis. 

In the majority of cases, the polypide appears capable of retracting to about the level of the 
highest part of its associated thickened walls. This suggests that a degree of protection may be 
afforded by the thickening, and has some analogy to the situation found in other, calcified, 
bryozoans. It is not clear whether the mineral salts reported to be found in the body walls of 
Amathia (Ryland 1970) are associated with any particular feature, such as this autozooidal 
thickening. 

From the autozooidal organisation evident, there are indications that some further analogy may 
be drawn between species of Amathia and other bryozoans, in terms of colony integration and 
co-ordinated behaviour. Together with regular budding patterns and specific orientations of auto- 
zooids, the localised autozooidal thickening carries with it implications for the achievement of 
lophophore eversion (and retraction). The thickened areas of cuticle might resist the deformation 
required by the autozooid to change its volume and effect these actions. There is little constraint on 
independent action of autozooids in those groups with inner wall thickening; the outer face of each 
autozooid is able to move freely in response to the volume changes necessary. In groups with outer 
wall thickening, the implied compliant boundaries for each autozooid are those walls contiguous 
with other autozooids. Thus, attempted changes in the colume of any one autozooid might impinge 
on the status of those adjacent. If these adjacent autozooids resist a change, then the eversion in the 
original autozooid will be hindered. It may be inferred, therefore, that in some species with outer 
wall thickening, feeding may be a group activity. Advantages of group feeding would lie in 
combined feeding currents, enhanced by specific autozooid orientations within colony bounded 
space, (Winston 1979, McKinney 1984). Independent autozooid behaviour is more likely if: the 
thickened outer wall has localised weak patches acting as diaphragms; the wall is sufficiently folded 
to allow concertina-like accommodation of volume change; the thickening differential is low; 
there are co-ordinated inverse volume changes of autozooid pairs. It cannot be discounted, how- 
ever, that collective feeding may occur in either wall-type grouping, simply by co-operation of 
autozooids. Confirmation of possible patterns of feeding behaviour, however, requires the 
observation of living colonies. 

In the autozooid groups, no pairing of autozooids may be confidently assigned throughout a 
colony in any species (see page 309). Although the concept of biserial rows loses some ground, it 
cannot be discounted completely. There is thus equal possibility that the arrangement of auto- 
zooids into groups may have evolved in any of three ways: by unification of two separate single 
rows of autozooids with subsequent modifications; by the linear organisation of randomly 
clumped autozooids; by spatial condensing, with alternate displacement, of one single row of 
autozooids. All three hypothetical initial conditions have some analogues in extant ctenostomes; 
the first in Zoobotryon, the last two in species of Bowerbankia. Tenuous indications for origins via 
the third category may be inferred from the order of autozooid production on stolons. Autozooids 
in a group are developed in distal sequence, often making their appearance laterally displaced on 
alternate sides. It is possible, however, that this simply reflects the fact that growth proceeds 
distally through a sequence of interlocked autozooids, as autozooids may also be seen to be 
produced as equal pairs. 

As with many colonial organisms, a large epifauna is frequently associated with colonies of 
Amathia, presumably deriving benefit from the microenvironment of the colony interiors (see 
below). The colonies serve as a substratum for some organisms and as shelter for others. Great 
numbers of other bryozoans, coelenterates, crustaceans, annelids, algae, foraminifera and 



340 P. J. CHIMONIDES 

molluscs, are often found. In this context, the record of Amathia body walls containing calcium 
salts (Ryland 1970) needs re-investigation from material in which the absence of any encrusting 
calcareous epibionts is ensured, as these can be extremely diaphanous. It is not known if any of the 
associations are species-specific, or what other levels of interdependence may occur. The ecological 
criteria which determines distribution and survival of the species of Amathia are known in only 
most general terms, and nothing is known of the relative ecological requirements which epibionts 
and 'hosts' may have. All that might have been expected is that numbers of epibionts might be 
related to some simple factor, such as the degree of shelter a colony provides. However, Murray 
(1970) reported that the entire life-cycle of the gastropod Marginella minutissima is spent with 
Australian A. biseriata. In this case, the Amathia colony serves both as food substrate as well as the 
physical substratum. Murray's concluding suggestion was that it is the occurrence of the bryozoan 
which actually determines the mollusc's distribution. 

The observable specific variation, and the limited numbers of recognisable characters perceived 
in these non-rigid animals, has made past workers, for example MacGillivray (1895), Hastings 
(1927), d'Hondt (1979, 1983), variably reluctant to accept the existence of certain species. As a 
result it has been suggested that some species, for example A. lendigera and A. distorts, are almost 
ubiquitous. Wide geographic distributions, continuous or discontinuous, are not unknown 
amongst marine animals (Ekman 1967, Cook and Lagaaij 1973), and the genus has been reported 
from nearly all marine regions except the polar and subpolar seas. However, there is no evidence 
that any species of Amathia has ever achieved and maintained a cosmopolitan distribution. Any 
indications to the contary seem based on misidentifications. The problem is compounded in one 
instance; for two specimens, A. wilsoni and A. semiconvoluta ex Lamouroux collection (BMNH 
1899.7.1.6606 parts), there is doubt that the locality data and specimens actually belong together 
(see page 335). 

Although Rao and Ganapati (1975) reported 'Amathia distans 1 as 'an important fouling species 
at the Visakhaptnam Harbour', species of Amathia are not noted as fouling the hulls of sea going 
vessels, and there is no indication that shipping has any effect (cf. Ryland 1970) on distribution. 
From the information available (albeit that this reflects the situation around the turn of the 
century, when many of the specimens studied were collected) the species determined appear to have 
distributions which reflect modern oceanic current flows (see below). This is not unexpected, as 
Amathia colonies are sessile, and the geographic distribution of species would be greatly dependent 
on dispersal of colony fragments and larvae by water currents. 

Taken simplistically, the maintenance of widespread distributions suggests the need for 
adequate gene flow to help preserve the biological unity of each species (Sheppard 1975, Speiss 
1977), and may be influenced by physical criteria. To some extent, this would involve the effects of 
sperm dispersal. Assuming some general similarity of ctenostomes with other Bryozoa, the free- 
swimming life of the lecithotrophic larvae (Barrois 1877, Nielsen 1971, Zimmer and Woollacott 
1977), might be estimated at about 24 hours. Records of lecithotrophic larval life in Cheilostomata 
range from 20-75 minutes as in Parmularia (Cook and Chimonides 1985), to a maximum of 3-5 
days as in Crassimarginatellafalcata (Cook 1985). Under the same assumption of similarity, sperm 
life might be estimated as up to 1 hour (Marcus 1926 for Electra pilosa, Silen 1966 for Electro 
posidoniae). Lecithotrophic larval life in Bryozoa is generally held to be short and dispersal limited 
(Ryland 1976, Farmer 1977, Hay ward and Cook 1983). Similarly, the contribution sperm dispersal 
makes towards preventing speciation must also be limited. 

It is difficult to assess what contribution fragmentation makes towards species distribution; for 
the present, it is possible only to speculate on the effects of the factors involved. It is unlikely that 
colonies of Amathia would be susceptible to the same shear forces that might cause rigid, calcified 
colonies to fail structurally (Cheetham and Erikson 1983). The shape of Amathia colonies results 
partly from the exoskeletal function of locally thickened cuticles, but derives mainly from turgor 
pressure of the various coelomic fluids acting on the cuticles. The cuticles are flexible but non- 
elastic. Such an essentially hydrostatic support system would be capable of a great deal of deform- 
ation with subsequent recovery. Structural failure results when drag forces exceed tensile strength. 
Tensile strength of alcohol preserved specimens examined appeared subjectively high. The failure, 



AMATHIA 341 

near a bifurcation, of single stolons taken from distal tips of a specimen of A. brongniartii from 
Victoria Australia (BMNH 1984.12.4.1), was recorded at 80 grams. 

Additional resistance to fragmentation is likely in colonies with dense branching. In these, water 
flow effects are prevented from acting directly on all the constituent components, and the effective 
drag of a colony is less than expected (Cheetham and Erikson 1983). Under this condition, much 
water flow would be redirected around the colony, and this would place some emphasis on the 
external hydrodynamic profile that a colony presents. A possible reaction to this is suggested in the 
fact that autozooids are often arranged to face into the relatively sheltered space within the colony 
interior, as for example in A. wilsoni, A. woodsii, A.populea and A. guernseii. 

The characteristics of flexibility, reasonable tensile strength and hydrodynamic reaction are, 
however, the very features which have allowed Amathia species to spread into the kind of high 
energy environments, for example, much of southern Australia (Thomas and Shepherd 1982, King 
and Shepherd 1982) where, if only under severe storm conditions, fragmentation of colonies 
themselves must occur. In less extreme circumstances, for some species, fragmentation of the 
possible algal substratum might occur, setting entire colonies adrift. 

The longevity of adult colony pieces, is potentially much greater than that of the larvae and 
sperm. Under laboratory conditions at the BMNH, specimens of Flustrellidra hispida survived for 
over 6 months without their original algal substratum, which had rotted away. The colonies 
adopted a highly mishaped globular form, approximately 1-5 cms. maximum dimension, lying free 
on the gravel filter bed of their container. These colonies could be bowled around by very mild 
water movement, while the great majority of the autozooids forming their surfaces, retained the 
ability to feed. 

As colony fragments of Amathia do not readily float, it is to be expected that they will be 
transported well only whilst they are kept clear of the sea floor. Transportation and being kept 
clear of the sea floor will take place only as long as there is the appropriate energy in the water 
currents. More distant dispersal is possible if rafting on a more bouyant substratum, such as algae, 
occurs (Cheetham 1966, Cook and Lagaaij 1973). The success of any dispersals would require the 
eventual deposition of species in some suitable environment. Three levels of failure seem possible: 
that destination environments outside the recorded distribution are unsuitable (in which case, 
under certain circumstances, it is not impossible that remnants of at least some of these failures 
might be found); that dispersals do not reach wider transportation currents; that dispersals do 
reach wider transportation currents but suffer mortality en route, through loss of the 'raft' as the 
alga dies and rots. A. lendigera and A.pruvotican be algal epibionts. These species, if any, would be 
expected to have achieved very wide distributions, but this does not appear to be the case. Their 
distributions instead appear similar to those of well documented Lusitanian faunas (Hardy 1959, 
Ekman 1967, Tait 1986, Currie 1983) (cf. A. semiconvoluta recorded from the west coast of Africa 
to the Mediterranean). 

Regardless of the dispersal method of fragmentation products, direct survival of fragments 
would mainly be favoured by a low energy environment. Higher energy environments might allow 
survival only through subsequent release of larvae and their settlement. No colonies have been 
encountered where direct re-establishment of fragments is recognised to have occurred. It is quite 
possible, however, that colony fragments of variable size may re-attach and grow, and even that 
arborescent colonies resume their posture and growth form with the aid of rhizoids, in a process 
analogous to that observed in Parmularia (Cook and Chimonides 1985). However, whatever the 
frequency of fragmentation and outcome of subsequent events, the effects on distribution appear, 
for the present, to be of little significance. 

The earliest record of fossil Amathia is from the Late Cretaceous, with a species appearing in the 
Maastrichtian of The Netherlands (Voigt 1972, Cheetham and Cook 1983). The genus is not 
associated with very deep water, the deepest record encountered being 150 fathoms (275 metres 
approx.) for specimens collected off Bahia during the Challenger Expedition. It seems likely, 
therefore, that the genus achieved its present day tropical to cold-temperate distribution via shelf 
waters through Tethys and the Tehuantepec Channel, and to have traversed these regions before 
their closure in the mid Miocene (Ekman 1967, Cook and Lagaaij 1973, Haq 1981). It is obvious 



342 P. J. CHIMONIDES 

that more evidence is required to support these suggestions, although this may not be readily 
available, as non-boring ctenostomes have a poor preservation record (Cheetham and Cook 1 983). 

It is interesting to note that some of the species recognised, A. pinnata, A. woodsii, A. biseriata, 
seem to have been recorded exclusively from both south Africa and southern Australia. Parallels 
exist for other bryozoan species (Hayward and Cook 1983). This distribution is almost certainly 
the resultant of palaeogeographic factors rather than of modern current flows (see below), and 
implies that the genus was established and speciated by the time Africa has moved into relative 
isolation from its Antarctic association. This does not extend the theoretical age of the group much 
beyond the Maastrichtian however (see above). 

Although the imprecision of past records is criticized, the interpretation here of both A. 
brongniartii and A. pinnata from Australia, as two single species, rather than as species complexes, 
is prehaps lenient even on present evidence. Similarly, the specific genetic unity implied in each case 
for A. biseriata, A. woodsii and A. pinnata in both south Africa and southern Australia, although 
accepted here, must be viewed with caution. No linking distributions are recorded and gene flow 
through dispersal of sperm, larvae and colony fragments is not favoured over such distances and 
locations, and would not prevent divergence from occurring. Additionally, long term genetic 
stability of species is implied. 

In general, it may be said that the members of the genus have had time to become distributed 
widely. There has also been enough time for the effects of isolation and isolating mechanisms in 
demes to have come into play (Schopf 1977, Speiss 1977). Furthermore, if the cryptic speciation 
indicated by Thorpe and Ryland (1979), for species of the ctenostome Alcyonidium, has any 
parallel in this ctenostome group, further subdivisions within many of the groupings proposed here 
should be expected. 



Acknowledgements 

I would like to thank the following: Dr P. E. Bock as a Research Associate of the National Museum 
of Victoria; Dr A. H. Cheetham of the Smithsonian Institution; Dr J.-L. d'Hondt of the LBIMM at 
the Paris Museum and Dr C. Fransen of Rijksmuseum, Leiden, for information and the loan of 
specimens. I am grateful to: Dr R. V. Melville of I.C.Z.N. for advice; Dr P. J. Hayward, Swansea 
University; Dr J. D. Bishop and Dr N. J. Evans of the BMNH, for constructive criticisms; Miss 
B. C. Househam, Mr A. Ritch and Mr M. Viney formerly of the BMNH for their shouldering of 
distractive duties; the BMNH Photo Unit for photographs. I am grateful to Miss P. L. Cook for her 
support and encouragement in the execution of this work. 



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AMATHIA 345 

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Manuscript accepted for publication 19 September 1986 



346 



P. J. CHIMONIDES 



tp 







po 



Fig. 1 Schematic representation of erect colony components with reference orientations: (A) az 
autozooecia, paz proximal-most autozooid, rh rhizoid, s stolon, tp terminal process; (B.) interior wall 
thickening; (C.) exterior wall thickening; (D, E.) a anterior, d distal, / left lateral, p proximal, po 
posterior, r right lateral. 



AMATHIA 



347 






Fig. 2 Relative orientations of autozooecia and rhizoids about the stolon: (A.) A. guernseii; (B.) 

A.populea; (C.) A. woodsii; (D.) A. tricornis. 



348 



P. J. CHIMONIDES 





Fig. 3 Relative orientations of autozooecia and rhizoids about the stolon: (A.) A. lamourouxi; (B.) 

A.pinnata; (C.) A. plumosa; (D.) A. obliqua. 



AMATHIA 



349 







Fig. 4 Relative orientations of autozooecia and rhizoids about the stolon: (A.) A. brongniartii; (B.) 

A. biseriata; (C.) A. alternata; (D.) A. wilsoni. 



350 



P. J. CHIMONIDES 






Fig. 5 (A.) Relative orientations of autozooecia and rhizoids about the stolon in A. semiconvoluta. 
Exemplified by A. lendigera: (B.) normal autozooid arrangement on a triad of maternal and daughter 
stolons with sister stolons carrying autozooid displacements to each other; (C.) alternative autozooid 
arrangement on a triad of maternal and daughter stolons, the daughter stolons carrying identical 
displacements, both opposite to the condition on the maternal stolon, dl left daughter stolon, dr right 
daughter stolon, m maternal stolon. 



AMATHIA 



351 




Fig. 6 (A.) A. lendigera BMNH 1942.8.6. 1 5, Neotype, Chichester Harbour, UK. x 1 8; (B.)A.guernseii 
BMNH 1898.5.17.189, Holotype, Guernsey, Guernsey, UK x 17; (C.) A. intermedis BMNH 
1887.5.2.18, Holotype, Hastings, UK x 27; (D.) A. populea BMNH 1899.7.1.526, Lectotype, Natal, 
South Africa, site of rhizoid origin arrowed x 44. 



352 



P. J. CHIMONIDES 




Fig. 7 (A.) /4. lendigera BMNH 1942.8.6.15, Neotype, Chichester Harbour, UK x 8; (B.) A. guernseii 
BMNH 1898.5.17.189, Holotype, Guernsey, UK x 8; (C.) A. intermedis BMNH 1842.12.9.14, Belfast 
Bay, N. Ireland x 6; (D.) A.populea BMNH 1899.7.1.526, Lectotype, Natal, South Africa x 8. 



AMATHIA 



353 




Fig. 8 (A.) A. lendigera BMNH 1942.8.6.15, Neotype, Chichester Harbour, UK, palmate processes 
x 28- (B.) A. obliqua NMV H493 (65391) Syntype, Port Phillip Heads, Aus. x 10; (C.) A. lamourouxi 
BMNH 1887.12.10.70, Neotype, Port Phillip, Aus. x 20; (D.) A. obliquaNMV H493 (65391) Syntype, 
Port Phillip Heads, Aus., site of rhizoid origin arrowed x 57. 



354 



P. J. CHIMONIDES 




Fig. 9 (A.) A. lamourouxi BMNH 1887.12.10.70, Neotype, Port Phillip, Aus. x 8; (B.) A. cornuta sensu 
d'Hondt (A. woodsii) LBIMM 2821 part, TOcean asiatique', bifurcate terminal process arrowed x 8; 
(C.) A. lamourouxi BMNH 1899.7.1.3, New Zealand, apparent alternate branching x7; (D.) A. 
woodsii BMNH 1 883. 1 1 .29.27, Neotype, Port Jackson, Aus., rhizoid origin arrowed x 1 3. 



AMATHIA 



355 




Fig. 10 (A/M./wmataBMNH 1888.5.17.8 A, Lectotype, Port Phillip, Aus. x6;(B.)A.pinnataBMNH 
1888.5.17.8 C, Port Phillip Aus. x 8; (C.) A. wilsoni BMNH 1888.5.17.7, Syntype, Port Phillip, Aus. 
x 5; (D.) A. wilsoni BMNH 1888.5.17.7, Syntype, Port Phillip, Aus., rhizoid origin arrowed x 1 1. 



356 



P. J. CHIMONIDES 




Fig. 11 (A.) A. brongniartii BMNH 1888.5.17.6, Neotype, Port Phillip, Aus., rhizoid origin arrowed 
x20; (B.) A. brongniartii BMNH 1888.5.17.6, Neotype, Port Phillip, Aus. x8; (C.) A. biseriata 
BMNH 1887.12.10.90, Neotype, Port Phillip, Aus. x8; (D.) A. biseriata BMNH 1887.12.10.90, 
Neotype, Port Phillip, Aus., rhizoid origin arrowed x 20. 



AMATHIA 



357 




Fig. 12 (A.) A. plumosa NMV H494, Holotype, Port Phillip Heads, Aus. x 10; (B.) A. plumosa 
BMNH 1963.2.12.354, Western Australia, rhizoid origin arrowed x 18; (C.) A. tricornis BMNH 
1899.7.1.6600, Holotype, Australia, rhizoid origin arrowed x 14; (D.) A. convoluta (A. crispd) 
BMNH 1899.7.1.6607, Australasia x 10. 



358 



P. J. CHIMONIDES 




Fig. 13 (A.) A. alternata USNM 6307, Neotype, Cape Fear, N.C. USA x 4 (B.) A. alternata BMNH 
1964.7. 10.1A, New River Inlet, N.C., USA, showing polyrhizoid condition, the rhizoid origins indi- 
cated x 10; (C.)A.semiconvoluta BMNH 1912. 12.2 1.687, Adriatic, rhizoid origin arrowed x6;(D.)A. 
pruvoti LBIMM Bry 8205, Type x 8. 



British Museum (Natural History) 

The birds of Mount Nimba, Liberia 

Peter R. Colston & Kai Curry-Lindahl 



For evolution and speciation of animals Mount Nimba in Liberia, Guinea and the Ivory Coast is 
a key area in Africa representing for biologists what the Abu Simbel site in Egypt signified for 
archaeologists. No less than about 200 species of animals are endemic to Mount Nimba. Yet, this 
mountain massif, entirely located within the rain-forest biome, is rapidly being destroyed by 
human exploitation. 

This book is the first major work on the birds of Mount Nimba and surrounding lowland 
rain-forests. During 20 years (1962-1982) of research at the Nimba Research Laboratory in 
Grassfield (Liberia), located at the foot of Mount Nimba, scientists from three continents have 
studied the birds. In this way Mount Nimba has become the ornithologically most thoroughly 
explored lowland rain-forest area of Africa. 

The book offers a comprehensive synthesis of information on the avifauna of Mount Nimba 
and its ecological setting. During the 20 years period of biological investigations at Nimba this in 
1 962 intact area was gradually opened up by man with far-reaching environmental consequences 
for the rain-forest habitats and profound effects on the birds. Therefore, the book provides not 
only a source of reference material on the systematics, physiology, ecology and biology of the 
birds of Mount Nimba and the African rain-forest, but also data on biogeography in the African 
context as well as conservation problems. Also behaviour and migration are discussed. At 
Nimba a number of migrants from Europe and/or Asia meet Afrotropical migratory and 
sedentary birds. 

Professor Kai Curry-Lindahl has served as Chairman of the Nimba Research Laboratory and 
Committee since its inception in 1962. Peter Colston is from the Subdepartment of Ornithology, 
British Museum (Natural History), Tring, and Malcolm Coe is from the Animal Ecology 
Research Group, Department of Zoology, Oxford. 

1986, 129pp. Hardback. 565 00982 6 17.50. 



Titles to be published in Volume 52 



Miscellanea 

A revision of the Suctoria (Ciliophora, Kinetofragminophora) 5. The Paracineta 
and Corynophora problem. By Colin R. Curds 

Notes on spiders of the family Salticidae 1. The genera Spartaeus, Mintonia and 
Taraxetta. By F. R. Wanless 

Mites of the genus Holoparasitus Oudemans, 1936 (Mesostigmata: Parasitidae) in 
the British Isles. By K. H. Hyatt 

The phylogenetic position of the Yugoslavian cyprinid fish genus Aulopyge Heckel, 
1841, with an appraisal of the genus Barbus Cuvier & Cloquet, 1816 and the 
subfamily Cyprininae. By Gordon J. Howes 

Revision of the genera Acineria, Trimyema, and Trochiliopsis (Protozoa, 
Ciliophora). By H. Augustin, W. Foissner & H. Adam 

The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) with a 
systematic review, a synopsis of Pipistrellus and Eptesicus, and the descriptions of 
a new genus and subgenus. By J. E. Hill & D. L. Harrison 

Notes on some species of the genus Amathia (Bryozoa, Ctenostomata). 

By P. J. Chimonides 



Printed in Greal Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 



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