BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE, BIOLOGIE, 71-S UPPL.: 7-I6, 200 I BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN, BIOLOGlE, 71-SUPPL.: 7-16, 200 I

A new myosoricine shrew from the Congo Basin referable to the forgotten genus Congosorex (Mammalia: Soricidae) by R. HUTTERER, P. BARRIERE & M. COLYN

Abstract DI PPENAAR, 1978), but in principal remained a synonym of what was fonnerly regarded as genus Myosorex. HUTTERER We report on the discovery of a small, short-tailed species of ( 1993, 1995), after having examined the holotype specimen shrew in lowland forest of the western Congo Basin, which in the Tervuren Museum, raised Congosorex again to generic resembles the enigmatic Congosorex polli, known from a level. However, as the type species C. polli was never col­ si ngle specimen collected in 1955 south of the Congo River. lected again, the genus remained an enigma. A new population, represented by 15 specimens, is referred to a new species of that genus. Di fferences to Myosorex and In 1995 and 1996, faunal surveys were conducted in three different forest sites in the Republic of Congo (RC) and in the Surdisorex, two other African myosoricine genera, are dis­ Central African Republic (CAR) by the two junior authors of cussed. Congosorex is now known from fo ur localities in the the University of Rennes 1 for the ECOFAC project. During central African forest black. The two species are separated by these surveys, 15 small and short-tailed shrews were ob­ the Congo River. tained which at close inspection turned out Lo belong to the Key-words: Insectivora, Soricidae, Congosorex, systemat­ long-forgotten genus Congosorex. Fourty-five years after its ics, biodiversity, Africa. description the genus is now re-discovered, but the species turned out to be different from the type species of the genus, Congosorex polli. The present paper serves to define the new Résumé species and to discuss its biogeographical and systematical significance. Nous rapportons la découverte, en fôret de plaine du bassin du Congo occidental, d'une petite espèce de musaraigne à queue courte rappelant J' eni gmatique Congosorex polli connu par un unique spé­ Material and methods cimen coll ecté en 1955 au sud du fl euve Congo. Une nouvelle popu­ lation, représentée par 15 spécimens, est attribuée à une nouvelle 15 specimens of the new shrew were collected in pitfall traps espèce de ce genre. Nous discutons de ses différences avec during inventory studies in the CAR and in the RC. Speci­ Myosorex et Surdisorex, deux autres genres de myosoricinés afri­ cains. Congosorex est désormais connu de quatre localités du bloc mens were weighed in the fie ld and subsequently preserved foresti er d'Afrique centrale, les deux espèces étant géographique­ in tluid. Voucher specimens form part of the collections cur­ ment séparées, l' une de l'autre, par Je fl euve Congo. rently stored at the Station Biologique de Paimpont, Université de Rennes l. Other specimens mentioned in this Mots-clés: lnsectivora, Soricidae, Congosorex, systématique, report fonn part of the collections of the following institu­ biodiversité, Afrique. tions: Musée Royal de l' Afrique Central, Tervuren (MRAC), Museum National d' Histoire Naturelle, Paris (MNHN), Introduction Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn (ZFMK), American Museum of Natural His­ tory, New York (AMNH), Carnegie Museum ofNatural His­ The African shrew Congosorex polli (HE IM DE BALS AC & tory, Pittsburgh (CM). LAMOTTE, 1956) is one of the least known mammals of the world, being the onl y representative of its genus, and known External and cranial measurements presented in this study by only one specinien, the holotype. This unique animal was (Tab. 3) were taken by RH from the preserved fluid speci­ collected by Stuart A. MARKS in 1955 at Lubondaie [near mens and the dry skulls using an electronic caliper. Ali meas­ Tshimbulu] in Kasai province, southern Democratic Repub­ urements are given in millimetres (mm) and body masses in lic of Congo (DRC), and subsequently named Myosorex grams (g). Names of authors and publication dates of species (Congosorex) polli by HEIM DE BALSAC & LAMOTTE ( 1956). discussed in the text are listed in HUTTERER ( 1993, 1995) and The new subgenus was occasionall y (HEIM DE BALSAC, WOLSA N & HUTTERER ( 1998) where complete citations o f 1967) treated as a fu ll genus or ubgenus (MEESTER & species and type localities can be fou nd . 8 R. HUTTERER, P. BARRIERE & M. COLYN

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Fig. 1. - Myosoricine crown pattern of the lower p4 in (from Ieft to right) Myosorex lo11gica11dat11s (ZFMK 81. 1293), Congosorex ve rheyeni n. sp. (R 16854), and Surdisorex norae (A MNH 87262).

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Fig. 2. - Sketches of head and ri ght fo refoot of Myoso rex varius (CM 94953, upper left figure), Congosorex verheyeni n. sp. (R 18344, upper right figu re), and Surdisorex po/11/u s (CM 2841, bottom); based on study skin s and fluid-preserved specimens. A new myosoricine shrew from the Congo Basin referable to the forgotten genus Congosorex 9

What is a myosoricine shrew? Table 2. Relat ive tail length RTL(% of head and body length) and condyloincisi ve length CIL (mm) in myosoricine shrews of Africa, based on the respecti ve Sorne confusion exists in the literature about the correct taxo­ holotype speci mens where available. nomie assignment of Myosorex and related fom1s. Here we place Myosorex; Congosorex, and Surdisorex provisionally Taxon RTL CIL in the tribe Myosoricini KRETZOI , 1965, of which Myosorex Surdisorex 11 orae 23. l 26.5 is the type genus (KRETZOI, 1965). It has long been known that this genus shows some ancestral characters (HEIM DE Surdisorex po/11/us 32.6 24.5 BALSAC, 1966, 1967). A biochemical study by MADDALE NA Co 11 gosorex 1·erheye11i n. sp. 30.2 17.8 & BRONNER ( 1992) showed that Myosorex is not closely re­ Co11go.wrex polli 40.0 20.3 lated to either Soricini/ae or Crocidurini/ae, the latter being Myoso rex ru111pii 41.7 22.7 the taxon in which Myosorex was most often pl aced. Al­ though KRETZOl 's name is available, as pointed out for ex­ Myoso rex kilwulei 47.0 20.0 ample by JAM MOT ( 1983) and HUTTERER (1993, 1995), ex­ Myoso rex babaulti 48.0 21.7 tant Myosorex were recently regarded by some authors Myoso rex cc~/er 50.0* 23.2* (MADDALENA & BRONNER , 1992; BEDFORD, BERN ARD & BAXTER, 1998) as members of the Crocidosoricinae, a taxon Myoso rex rarius 50.6* 22.4* named by REUMER (1987) to cover some extinct Miocene Myoso rex geara 53.3 21.0 1 shrews of Europe. The phylogenetic relationships between Myosorex sc/ateri 53 .5 25.2 Crocidosoricinae and extant Myosorex have never been in­ Myoso rex eise11tra 111i 56.3 22 .6 vestigated, but even if investigations demonstrate a closer relationship between the two groups the name Myosorici ni Myoso rex blarina 56.8 22.1 would have fo rm ai priority. QUÉROU IL et al. (2001) studi ed Myosorex okuensis 58.3 21.9 165 rRNA seguences of Congosorex (samples of the new Myosorex te11 11is 59.2 21.7 species described herein) and Myosorex and their results .show that both ge nera are sister taxa and belon g to a lineage Myosorex zi11ki 64.0 23.2 which is ancestral to Crocidura, Suncus, Sylvisorex, Myo.w rex /011 gica 11datt1s 75 .3 22 .9 Scutisorex, Ruwenza risorex and Paracrocidura. The ta xo n Myoso rex schalleri 83 .8 18.9 Myosoricini there fo re certainly includes Con.gosorex and Myosorex, while the inclusion of Surdisorex is yet based on *no types ava il able, means taken from MEESTER & DIPPENAA R 1978 morphological characters alone. The key character for the defi nition of the Myosoricini (JAMMOT, 1983) and Crocidosori cinae (REUMER, 1987, 1998) was the shape of the Externally, the long-clawed forefoot and the reduction of the fourth lower premolar (Fig. 1) : the V-shaped pattern of the ear conch separates Surdisorex from the other two ge nera. occlusal surface occurs in ail three extant genera, and is also The differences between Myosorex and Congosorex are more shared by Ca1posorex, Clapasorex, Crocidosorex, Florinia, subtle: Congosorex has a minute eye opening, and the scales Lartetium, Miosorex, and a number of other fossil taxa on the forefeet are large versus small in Myosorex. The tail (RE UMER, 1998). However, the phylogeneti c relationships length also separates the tlu·ee taxa: relative tai l length (Tab. between the fossil and extant taxa have not been studied 2) is shortest in Surdisorex (23-33%), medium in Congosorex properl y, and the inclusion of ail taxa sharin g a single tooth (30-40%), and long in Myosorex (42-84%). The ex\ant character into one clade is hi ghly hypotheti cal. Myosori cini of Africa contain 18 species: 14 species of MEESTER (1953) convi ncingly di scussed the generic di stin c­ Myosorex (STAN LEY & HUTTERER, 2000), two species of tion of Myosorex and Surdisorex, while Congosorex has only Con.gosorex (this study), and two species of Surdisorex recentl y been recognized (HUTTERER, 1993, 1995). Tables 1 (HEIM DE BALSAC & MEESTER, 1977). The fossi l record of and 2 present some characters which define the three taxa. the group is poor: onl y Myosorex is doc urnented frorn Miocene to Holoceni;: sites in Africa (BUTLER & GREEN­ WOOD, 1979; BUTLER , 1998). Tab le 1. Some distingui shing characters of the three African myoso ri cine genera. " Systematic descriptions Character Myosorex Congosorex Surdisorex Tai! medium to long short short Family Soricidae G. FISCHER, 18 17 Claws of forefoo t short short long Subfamily Soricinae G. FISCHER , 1817 Tribe Myosoricini KRETZO I, 1965 Scales on forefoo t small . large large Ear conch norm al normal/reduce d absent Genus Congosorex (HEIM DE BALSAC & LAMOTTE, 1956) Eye opening normal minute min ure Upper unicuspids 4 3 3 TYPE SPECIES Lowe r p3 present presen1/absent absent Myosorex polli HEIM DE BALS AC & LAMOTTE, 1956 10 R. HUTTERER, P. BARR IERE & M. COLYN

Emended diagnosis: Terrestrial shrews of the central African PARATYPES forest with reduced ear, eye, tait and limbs and a myosoricine 4 males, 1 female, field numbers R 16746, R 16764, R 16854, dentition. Tait 30-40% of head and body length; claws of R22903, R23014 (=MNHN CG 2000-467), ail collected forefoot of medium length. Upper dentition with 3 unicuspid 1996 at the same locality as the holotype. Ali specimens pre-' teeth and lower dentition with only one unicuspid tooth; p3 served in fluid , with most skull s extracted and cleaned. absent in most specimens. Additional cusplets present on the lingual side of upper un icuspids, lower p4 with V-shaped crown pattern. REFERRED MATERIAL 2 females. R 16825, R 16835, collected in 1996 by a team of DISTRIBUTION the University of Rennes 1 at the Great Escarpment (Ol .04N, l 4.29E), Parc National d' Odzala, Republic of Congo; 4 Confined to the Congo Basin, Africa. males, 3 females, Rl8126, Rl8283, Rl8298, Rl8299, Rl8321 , Rl8341 , Rl8344, collected 1995 by a team of the University of Rennes 1 at Bambio, Ngotto Forest (02.47N, Congosorex polli (HEIM DE BALSAC & LAMOTTE, 1956) i 6.25E), Central African Republic. Ali specimens preserved in fluid, with skull s extracted and cleaned. Tissues of several HOLOTYPE specimens are kept for genetical research, some of which have already been studied (QUÉROU IL et al. , 1998). MRAC 23.234, animal in spirit, skull extracted; collected by STUART A. MARKS 1955 at 'Lubondaie via Tshimbulu' (06.30S, 22.39E), Kasai province, DRC. ETYMOLOGY The species is named for our colleague and friend Walter DIAGNOSIS Verheyen in recognition of his broad and stimulating re­ search on the systematics and biogeography of African mam­ « .. un Myosorex dont la denture a évolué selon la li gne mals. orthogénétique générale vers le type Surdisorex, mais qui n'a pas subi ! 'évoluti on secondaire de loreille et des extrémités de ce dernier» (HEIM DE BALSAC & LAMOTTE, 1956: 155).

Other characters mentioned in the original description are: Table 3. Body mass (g), external and cranial measurements (mm) of head large compared to the rest of the body; ears proportion­ Congoso rex po/li and C. 1·er/J eye11i n. sp. all y larger than in Myosorex varius and M. cafer; hindfeet short, forefeet not elongated as in Myosorex; tail extremely Character C. pu/li C. rerheyeni n. sp. short (24 mm); colour very dark; hair very short; dentition holotype holotype mean+SD, (n) remarkable for the absence of the P3 (as in Surdisorex) and ZFMK 99.932 the presence of a tin y p3 (as in Myosorex); m3 very large; for Extcrnal measurements see table 3. Body mass - 5 7.07+2.23 ( 14) Head and body length 60* 6 1.5 59.24+1.84 ( 15) DISTRIBUTION Tai! length 24 18.6 19.59+1.63 ( 15) Known only from the type locality (Fig. 7). Hindfoot c. u. 11. 2 10.2 10.34+0.45 (15)

Hindfoot s.u. 10 9.5 9.2 1+0.36 (15) REMARKS Ear length 6.5 5.7 6. 14+ 1.03 ( 12) Congosorex polli has not been collected again and thus re­ Cra niai main s known from the holotype only. By application of the IUCN Red List criteria, the species should be considered as Condylo-in cisive length 20.34 17 .77 18.44+0.38 (10) «extinct». We suspect, however, that a careful survey of suit­ Palatal length 8.0 1 6.94 7.72+0.40 ( 10) abl e habitats near the type locality will show that the species Interorbital width 436 4.37 4.34+0. l 7 ( 10) still exists. A faunal survey has not been conducted in that region since the work of Stuart A. MARKS. Zygo111atic breadth 6.3 1 6.33 6,34+0. l l ( 10) Greatest width 10.38 1002 9.98+0. !9 ( 10)

Congosorex ve rheyeni n. sp. Height of cranial capsule 5.20 4.86 5.06+0.24 (10) Postglenoid width 7 08 6.9 1 6.90+0 15 ( 10) HOLOTYPE Upper toothrow length 8. 74 7. 7 1 8. 10+0.23 (11 )

ZFMK 99.932, female specimen preserved in alcohol with Lower toothrow length 7.94 7.04 7.37+0.22 ( 10) the skull extracted and cleaned. Field number R 16761. Col­ Coronoid height ( Il ) lected on 19 October 1996 by members of the University of 5.24 4.79 4.86+0. 16 Renn es 1 and the ECOFAC team at Mbomo (00.24N, ''the 01i ginal label gives 55 mm. but RH 111 easured 60 111111 fro 111 thc l 4.44E), Parc Nati onal d' Odzala, RC. holotypc A new myosoricine shrew from the Congo Basin referable to the forgotten genus Congosorex 11

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Fig. 3. - Congosorex verh eyeni n. sp., freshly captured specimen photographed by Ch. RATIER in Janu ary 1997 at the Great Escarpment, Odzala National Park, Republi c of Congo. About twice natural size.

TYPE LOCALITY short and weak caudal vertebrae (Figs 5, 6); pale coloured Mbomo (00.24N, l4.44E), Parc National d'Odzala, RC. For and covered with short black hairs over its entire length. a map of the park , see V ANLEEUWE, CAJANI & GAUTHIER­ Skull (Figs 4a-b) short and stout; greatest width of skull half HION ( 1998). of its total length (Tab. 3). Snout portion of skull short, inter­ orbital constriction wide, and braincase large and strongly DIAGNOSIS angular. Upper and lower dentition principally as described for C. pol/i by HEIM DE BALSAC & LAMOTTE ( 1956), except Differs from Congosorex polli by smaller size, much shorter for size. Upper unicuspid teeth with additional lingual tait and fe et, and by shorter ears. Skull smaller and stouter, cusplets, upper M3 very large. Lower p4 with the typical V­ particularly the snout. Lower p3 abse nt in most specimens. pattern of a myosoricine shrew (Fig. l ). For measurements, see table 3. GEOGRAPHIC VARIATION DESCRIPTION No size or colour variation was noted arnong the samples A very small and stout shrew (Fig. 3) with a very short tait ; studi ed. In the Odzala popul ati on, two specirn ens had addi­ body proportions recall small spec ies of Blarina or C1)ptotis. tional teeth: R22903 had a tiny pair of additional unicuspids Tait length less than one third of combined head and body (P3) in the ma ill ary, and R230 14 showed a tiny p3 on the length (Ta b. 3). Colour aspect of body dark brown, with no labial side of the left rn andibl e. contrast between dorsum and venter. Body hairs short, Smm on dorsum and 3mm on venter. Head comparatively large; COMPARISON di stance from tip of nose to back of ear co nch about one third of head and body length . Nose and lips fl esh-coloured. Vi­ From the holotype of Congosorex polli the new species dif­ brissae sparse and short; longest vibrissa 12 mm . Eye open­ fers by much shorter lirnbs, and by a shorter ear (Tab. 3). The in g minute and hidden in the fur. Ear co nch present but re­ skull s are sirnilar in overall shape (Fig. 4), but the anteri or duced in size and covered by body hairs. Forefeet short with part of the skull is more compressed in C. verheyeni, the in ­ moderately developed digits and claws; scales coverin g the terorbital constricti on is shorter and wider, whereas the size dorsal surface of the manus large. Hindfoot also short with of the braincase is similar. This results in a much shorter short claws (Fig. 3) and large scales. Tait very short, with 11 toothrow and a shorter skull length of the new species. The 12 R. HUTTERER, P. BARRIERE & M. COLYN

A B c

Fig. 4A-C. - A, Congoso rex verh eyeni n. sp., (holotype ZFMK 99.932), skull in dorsal view; B, same specimen, ventral view (total length of skull 17.8 mm); C, Congosorex polli, holotype skull in ventral view (total length 20,34 mm, reproduced from H EIM DE B ALSAC & L AMOTIE 1956, fi g. 5).

A

B

Fi g. 5A,B. - A, X-ra y of a specimen of A, Myoso rex va ri 11 s (ZFM K 79.274) and B, Co 11 gosorex ve rh eye11.i n. sp. (R 1832 1); note the short tail and hind li mbs o f lhe new species. Sli ghtl y enl arged. A new myosoricine shrew from the Congo Basin referable to the forgotten genus Congosorex 13

B

Fig. 6A,B. - X-rays of the pelvic region, hind limbs and tail of A, Myosorex va ri us (ZFMK 79.274) and B, Congosorex verheyeni n. sp. (R 1832 1). P = pelvis, F = femur, T = tibi a; caudal vertebrae 1-5 numbered. tooth formul a differs between the two species in that the DISTRIBUTION lower p3 is absent in most skulls (20 out of 22 mandibles Known from three locali ties in the Republic of Congo and examined) of C. verheyeni, while it is present in the single one locality in the Central Afri can Republic (Fig. 7). Both specimen of C. polli. Unless further specimens of C. polli localities are within the western Co ngo basin (COLYN, 1999) will be found , it remains unclear whether this is the typical and on the 1~i g ht bank of the Congo River. condition in that species. If the absence of p3 will turn out to be a typical trait of Congosorex, then this character would be shared with Surdisorex, along with the reduction of P3, in ECO LOG Y contras! to Myosorex. A low degree of variation in the pres­ In the Parc National d'Odzala most specimens were caught ence/absence of P3 or p3 is know from Myosorex and in pitfalls placed in primary fo rest, and some were caught in Surdisorex as we ll (MEESTER, 1953, 1955), and probably re­ degraded primary Forest nexl to a road where palm lrees and fl ects the close relatjonship between these genera. other anthropogenic vegetation grew. In the Ngo tto Forest Compared Lo a typical Myosorex (Figs 5, 6), Co11gosorex (CAR), one C. verheyeni was caught in secondary Forest, and verheyeni is a stoutl y built animal with very short Jimbs. Ail further six in pri mary forest. In the Parc Nati onal d' Odzala, elements of the skeleton are reduced in length ; in the the new species occurs together wi th at least 15 other shrew hindlimb, fo r example, not onl y the digits of the foot are spècies. Almost ail of them are forest species. Sorne are short, but also the ti bia, femur and pelvis (Fig. 6). The short widely distributed in the lowland Forest of the Congo Basin tail is not onl y caused by a reducti on of the number of caudal ( Sylvisorex joh11sto11 i, Crocidu ra dolichura, C. littoralis, C. venebrae ( 11 versus 14 in Myosorex). but also by a shorten­ hildegwdeae ), and some are confined to the western part of ing of the vertebrae. the Congo Basin. Examples are S1111c 11s remyi. Sr lvisorex 14 R. HUTTERER, P. BARRIERE & M. COLYN

CJ Lowland tropical forest CJ Woodland/grassland with scattered forest patches

Fig. 7. - Localities where Congosorex verheyeni n. sp. ( 1, 2, 3) and C. polli (4) were found. ( 1) Ngotto fo rest, Central African Republic; (2) Great Escarpment, Republic of Congo; (3) Mbomo, Odzala National Park, Republic of Congo; (4) Lubondaie, Democratic Republic of Congo.

ollula, S. cf ko11gane11sis, Paracrocidura schoutede11i, cently the scattered records of shrews in the Congo Basin did Crocidura crenata, and C. grassei. A detailed study of the not show any biogeographical pattern . With increasing ecology of the shrew communities occurring at these locali­ knowledge patterns emerge, and will be more evident after ti es is in preparation by one of the authors (PB). ail the material assembled during the various inventories will have been identi fied and analyzed. In addition Co11gosorex verheye11i seems to have a very special ecology. Apart from Conclusions the present study, the species was not collected during previ­ ous inventories in the Congo Basin where the same method­ The discovery of a new species of shrew which shares most ology (pitfall trap lines with drift fences) were used. Such characters with the enigmatic Myosorex (Congosorex) polli, studies were conducted in central and northern Gabon supports the recognition of Congosorex as a separate taxo­ (BROSSET, 1988; GOODMAN, HUTTERER & NGNEGUEU, nomie unit. HEIM DE BALSAC & LAMOTTE (1956) regarded 2001), in western and southern Cameroon (HUTTERER & Congosorex as a transition form between Myosorex and SCHLITTER, 1996; COLYN, CORNELI US & PERPÉTE, 1996), Surdisorex, a view parti y supported by the set of characters and in Equatorial Guinea (LASSO, HUTTERER & RIAL, 1996). summarized in Table 1. Whether this idea reflects phylogeny Also the study of RAY & HUTTERER (1996), who analyzed wi ll ·be unknown unti l a larger set of species of Myosorex, remains of shrews from carnivore scats in Central African Surdisorex, and Congosorex has been studied in a Republic, fai led to record any Congosorex. Some specimens " . phylogenetic context. At present it seems best to treat all in our study were collected in «borderline habitat», such as three taxa as separate genera. secondary fores t, open fo rest with grass cover, and not in pri­ The distribution of Congosorex is interesting: ail known mary forest where most of the other studies were conducted. specimens of Congosorex verheyeni are from localities on The minute eyes, short ears and the short tail suggest that the ri ght bank of the Congo River, white the single specimen Congosorex verheyeni may be somehow adapted to a subter­ of C. polli was collected on the left bank of the Congo River ranean life, although the short claws tell that the species is (Fig. 7). Such a pattern was found also in other mammals, for not a specialized digger. Congosorex verheyeni is a most in­ ex ample in rodents of the genus Lophuromys ( L. teresting discovery among the insectivores of the Congo Ba­ nudicaudatus north and L. huttereri south of the Congo si n, which would deserve a careful biological and ecological River; VERHEYEN , COLYN & HULSELMANS , 1996). Until re- study in the future. A new myosori cine shrew from the Congo Basin referable to the forgotten genus Congosorex 15

Acknowledgements HEIM DE BALSAC, H., 1967. Faits nouveaux concernant les Myosorex (Soricidae) de l'Afrique orientale. Mammalia, 31:610- Field work in Africa was possible through the support of the 628. European Community Commission (ECOFAC «Conserva­ HEIM DE BALSAC, H. & LAMOTTE, M., 1956. Evolution et ti on et utilisation rationelle des ECOsystèmes Forestiers en phylogénie des Soricidés africains. Ma111111 alia, 20: 140-167. Afrique Centrale» managed by AGRECO, GEIE, BDPA, HEIM DE BALSAC, H. & MEESTER, J., 1977. Part 1: Order SCETAGRI, SECA, CIRADFORÊT) and BioFAC DGVIII lnsectivora. Pp. 1-29 in: «The mammals of Africa: an identification projects. Materi al support and help in the field were provided manual» (MEESTER, J. & SETZER, H.W. , eds.). Smithsonian Insti­ by the ECOFAC project leaders: Jean-Marc FROMENT (RC), tution Press, Washington, D. C. Alain MONTFORT and Alain FENELON (CAR) , and by HUTTERER, R., 1993. Order lnsectivora, Pp. 69-130, in: «Mammal Nicolas HULOT («Okavango» TV programme). We thank Species of the World, a taxonomie and geographic reference, also the Congolese and Central African authorities of the Second ed.» (WILSON, D.E. & REEDER, D.M., eds.). Smithsonian «Ministère des Eaux et Forêts» and «Ministère des Eaux, Institution Press, Washington, D.C. 1206 pp. Forêts, Chasses et Pêches et du Tourisme», respecti vely. The aid of the research assistants Olivier PERPÉTE (Université de HUTTERER, R. & SCHLITTER, D. A., 1996. Shrews of Koru p Na­ ti onal Park, Cameroon, with the description of a new Sylvisorex Rennes 1) , Gabriel ÜSSESSA (RC), Gabriel Y ANGOU NDJARA, (Mammalia: Soricidae). ln: «Contributions in Mammalogy: A Me­ Robert KORISTAN MARO and Denis SACOM (CAR) have mori al Volume Honoring Dr. J. KNOX JONES , jr. » (GENOWAYS, been indispensable throughout the field work. We are also H. H. & BAKER, R.J ., eds), pp, 57-66, Museum of Texas Tech Uni­ grateful to the institutions and curators who provided com­ ve rsi ty, Lubbock. parati ve material and working facilities: American Museum JAMMOT, D., 1983. Evolution des Soricidae. Insecti vora, Mamma­ of Natural History (Guy G. MUSSER), Carnegie Museum of lia. Symbioses, 156: 253-273. Natural History, Pittsburgh (Duane A. SCH LITTER, Suzanne MACLAREN), Tervuren (Wim VAN NEER). C. KURZ (lnstitute KR ETZOI, M., 1965. Drepanosorex - neu defini ert. Vertebrata of Palaeontology, Bonn) provided the X-rays; Ch. RATIER H1111 garica, 7: 117-129. allowed to reproduce his photograph of the new species, and LASSO, C. , HUTTERER, R. & RI AL, A., 1996. Records of shrews Gustav PETERS kindly checked the English text. (Soricidae) from Equatorial Guinea, especiall y fro m Monte Alen National Park. Ma111 111alia, 60: 69-76. MADDALENA , T. & BRONNER, G., 1992. Biochemical systematics Literature cited of the endemic African genus Myosorex GRAY, 1838 (Mammalia; Soricidae). Israel Journal of Zoo log)~ 38: 245-252. BEDFORD, J.M ., BERNARD, R.T.F. & BAXTER, R.M. , 1998. The 'hy­ MEESTER, J. , 1953. The genera of African shrews. Annals of the brid' character of the gametes and reproducti ve tracts of the African Transvaal Muse 11111, 22: 205-214. shrew, Myosorex varius, supports its classification in the Crocidosoricinae. Journal of Reproduction and Fer1ility, 11 2: 165- MEESTER, J., 1955. Fossi! shrews of South Africa. Annals of the 173. Transvaal Museum, 22: 271 -278. BROSSET, A., 1988. Le peuplement de mammi fères insectivores des MEESTER, J. & DIPPENAAR, N.J ., 1978. A new species of Myosorex fôrets du nord-est du Gabon. Revue d ' Ecologie (la Terre et la Vie), from Knys na, South Africa (Mammali a: Soricidae). Annals of the 43: 23-46. Transvaal Museum, 31: 29-42. BUTLER, P.M ., 1998. 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