NEW SPECIES OF PARAPHIOMYS (RODENTIA,

THRYONOMYIDAE) FROM THE LOWER MIOCENE

OF AS-SARRAR, SAUDI ARABIA

by RAQUEL LOPEZ ANTONANZAS and SEVKET SENUMR 5143 du CNRS, Departement Histoire de la Terre, Museum national d’Histoire naturelle, 8 rue Buffon, F-75005 Paris, France; e-mail: ralopan@mnhn.fr,

sen@mnhn.fr

Typescript received 1 July 2003; accepted in revised form 12 December 2003

Abstract: The family Thryonomyidae is represented in the

Lower Miocene of Saudi Arabia by a single species, Paraphio-

mys knolli sp. nov. This new taxon differs from all other

thryonomyids in being small, lower molars having a short

metalophulid II and an isolated anterolabial cuspid, and

upper molars being antero-posteriorly compressed and pen-

talophodont. A cladistic analysis involving all extinct and

extant species of thryonomyids is provided. Paraphiomys

knolli branches as the sister-species of Paraphiomys pigotti,

type species of the genus.

Key words: Rodentia, Thryonomyidae, Paraphiomys, phylo-

geny, Miocene, Saudi Arabia.

The Saudi–French palaeontological Research Project

was carried out between 1977 and the early 1980s and

resulted in the discovery of several Lower and Middle

Miocene mammal localities near the coast of the Ara-

bian Gulf, Saudi Arabia. Thomas et al. (1982) published

preliminary results of this field work and recognized at

least 66 species from the Early Miocene (tentatively cor-

related to MN 5) faunas of the As-Sarrar area (Text-

fig. 1). Four sites there have yielded rodent remains,

but representatives of the family Thryonomyidae are

recorded only at localities 8 and 21. Thomas et al.

(1982, p. 129) provisionally assigned this material to

Paraphiomys sp. They briefly described it as pentalopho-

dont with well-developed mesolophs and mesolophids.

Flynn et al. (1983) described and illustrated the right

lower molar AS8-1100, on the basis of which they

announced the presence of a plesiomorphic thryonom-

yid in the As-Sarrar fauna. Finally, Winkler (1992,

p. 244) considered this taxon to be possibly new, at the

species or even genus level.

The thryonomyid dental material collected in Saudi

Arabia is especially important from a palaeogeographical

point of view because it bridges a gap between African

and Pakistani representatives. The aim of this paper is to

present a detailed study of the thryonomyid dental mater-

ial that has been recorded from the Lower Miocene of the

Dam Formation in Saudi Arabia and to compare it with

all Oligocene and Miocene thryonomyid species known to

date.

The terminology for tooth morphology follows that of

Winkler (1992).

Abbreviations: AMNH, American Museum of Natural History,

New York, USA; AS, As-Sarrar; CGM, Geological Museum,

Cairo, Egypt; GSW, Geological Survey, Windhoek, Namibia;

H-GSP, Howard University-Geological Survey of Pakistan; IPS,

Instituto de Paleontologıa, Sabadell, Spain; KNM, National

Museums of Kenya, Nairobi, Kenya; MNHN, Museum national

d’Histoire naturelle, Paris, France; NMAA, National Museum,

Addis Ababa, Ethiopia; UM, Universite des Sciences et Tech-

niques du Languedoc, Montpellier, France.

SYSTEMATIC PALAEONTOLOGY

Order RODENTIA Bowdich, 1821

Family THRYONOMYIDAE Pocock, 1922

Genus PARAPHIOMYS Andrews, 1914

Type species. Paraphiomys pigotti Andrews, 1914

Paraphiomys knolli sp. nov.

Text-figure 2A–F

Derivation of name. In honour of the palaeontologist Fabien

Knoll.

Holotype. AS8-1100 (Text-fig. 2A), a right m1-m2.

[Palaeontology, Vol. 48, Part 2, 2005, pp. 223–233]

ª The Palaeontological Association 223

Paratypes. AS21-1101, left D4 (Text-fig. 2B); AS8-1102 (Text-

fig. 2C), AS8-1103 (Text-fig. 2D), right M1-2; AS8-1104 (Text-

fig. 2E), left M3; AS8-1105, right M3 (Text-fig. 2F). All material

is housed in the collections of the Laboratoire de Paleontologie

(MNHN); size data for each tooth are provided in Table 1.

Localities and type horizon. Loc. 8 (type locality) and Loc. 21,

As-Sarrar area, Saudi Arabia. Dam Formation; late Early Mio-

cene (c. 15–17 Ma).

Diagnosis. Species of Paraphiomys of small size with sem-

ibunodont teeth. Lower molars with three complete lophs,

plus a short metalophulid II, an isolated anterolabial cuspid,

and lacking anterolabial cingulum. Antero-posteriorly

compressed upper molars with five complete lophs and a

labially displaced hypocone.

Differential diagnosis. Paraphiomys knolli differs from

Gaudeamus aegyptius, Neosciuromys africanus, Apodecter

stromeri, P. orangeus, P. renelavocati, P. simonsi, P. ship-

mani, Kochalia geespei, Paraulacodus indicus, P. johanesi,

P. afarensis, P. roessneri, P. australis and the two living

species of Thryonomys notably in having metalophulid II

on the lower molars. It is distinct from Paraphiomys pig-

otti and P. occidentalis in being much smaller than these

and from Epiphiomys coryndoni in having a much shorter

metalophulid II. Paraphiomys knolli differs from P. hop-

woodi in being smaller, having the upper molars more an-

tero-posteriorly compressed and in showing a more

labially displaced hypocone. Finally, this species is distinct

from Sacaresia moyaeponsi in having the teeth semibuno-

dont whereas the latter species is hyperlophodont but

lower crowned with slender lophs.

Description

m1 or m2 (AS8-1100). The single lower molar from As-Sarrar

has a subrectangular occlusal outline. It has three transverse

lophs (metalophid, hypolophid and posterolophid) and a short

metalophulid II. This tooth shows an isolated and transversely

elongated anteroconid. Mesoflexid and metaflexid are approxi-

mately equal in length as well as in depth, and open lingually,

but the valley between the hypolophid and the metalophid is

wider than that between the hypolophid and the posterolophid.

The protoconid is situated slightly external to the hypoconid.

The hypoflexid is deeper than the mesoflexid and the metaflexid,

and it is almost transverse.

D4 (AS21-1101). A single well-preserved tooth (D4) has been

found at locality 21 of As-Sarrar. It is trapezoidal in occlusal

outline. The anterior half of the crown is narrower than the

posterior one. The tooth has an anteroloph, a protoloph, a meta-

loph, a posteroloph and a very short mesoloph. The protoloph

is incomplete and does not reach the labial side of the tooth.

The metaloph is well developed and connects lingually with the

posteroloph. The protoloph and the metaloph are anteriorly

directed. The hypoflexus is nearly transverse.

M1 or M2 (AS8-1102 and AS8-1103). Both specimens are badly

worn and lack their lingual portions. These teeth have an anter-

oloph, a protoloph, a metaloph, a posteroloph and a short,

transverse mesoloph. The anterior half of the crown appears to

be narrower than the posterior one. The paraflexus is longer and

slightly deeper than the metaflexus.

M3 (AS8-1104, AS8-1105). The third upper molars from

As-Sarrar are worn, but all their structures are easily recogniz-

able. The occlusal outline is rounded. The anterior half of the

crown is wider than the posterior one. These teeth have four

TABLE 1 . Measurements (mm) of the teeth of Paraphiomys

knolli sp. nov. from the Arabian Peninsula.

As-Sarrar Length Width

m1 or m2 AS8-1100 1Æ88 1Æ74

D4 AS21-1101 1Æ80 1Æ51

M1 or M2 AS8-1102 1Æ88 –

M1 or M2 AS8-1103 1Æ96 –

M3 AS8-1104 1Æ74 1Æ78

M3 AS8-1105 1Æ77 1Æ86

TEXT -F IG . 1 . Major African, Arabian, and southern Asian

Eocene–Miocene thryonomyid localities. 1, Thymiana (Chios

Island, Greece); 2, Beni Mellal (Morocco); 3, Jebel Zelten

(Libya); 4, Fayum (Egypt); 5, Ch’orora (Ethiopia); 6, Napak and

Bukwa (Uganda); 7, Lothagam and Samburu hills (northern

Kenya); 8, Baringo (central Kenya); 9, Songhor, Rusinga,

Maboko, and Fort Ternan (western Kenya); 10, Otavi

(Namibia); 11, Namib (Namibia); 12, As-Sarrar (Saudi Arabia);

13, Abu Dhabi (United Arab Emirates); 14, Sind Province

(Pakistan); 15, Zinda Pir Dome (Pakistan); 16, Potwar Plateau

(Pakistan). The grey area represents the present day distribution

of the family.

224 P A L A E O N T O L O G Y , V O L U M E 4 8

lophs (an anteroloph, a protoloph, a metaloph, a posteroloph)

and a short mesoloph. As shown by AS8-1105, the posteroloph

fuses with the metaloph during wear. In specimen AS8-1104, the

hypoflexus is oblique, pointing strongly anteriorly, whereas it is

nearly transverse in AS8-1105.

Comparisons

Gaudeamus aegyptius Wood, 1968. Gaudeamus aegyptius has

been recorded only from an Upper Eocene (?) horizon in the

Jebel el Qatrani Formation, Fayum Province, Egypt (Text-fig. 1).

The holotype of this species (CGM 26920) is a lower jaw with

d4-m2 and an unerupted premolar (Wood 1968, fig. 14C–E).

The lower molars of G. aegyptius mainly differ from Paraphiomys

knolli in lacking a metalophulid II and the anterolabial cuspid.

With regard to the upper molars, the two taxa are also morpho-

logically very different because of the absence of a mesoloph in

G. aegyptius.

Sacaresia moyaeponsi Hugueney and Adrover, 1991. The holo-

type (IPS-7660) of this species is a fragmentary maxilla with M1-2

from the Oligocene site of Paguera 1, Majorca, Spain. This species

is very different from Paraphiomys knolli. The lower molars have

four lophs whereas P. knolli has a basic morphology with three

lophs and a vestigial metalophulid II. S. moyaeponsi has an antero-

labial cingulum, but it lacks an anterolabial cusp. In contrast, the

lower molar of P. knolli shows the anterolabial cusp, but it lacks

the cingulum. Another difference between the two taxa is the

hyperlophodonty and slender lophs in all teeth of S. moyaeponsi

whereas the teeth of P. knolli are semibunodont. In AS8-1100, the

posterolophid curves towards the metaconid whereas it is nearly

straight and posteriorly directed in S. moyaeponsi.

The morphology of the upper molars of S. moyaeponsi is also

very different from that of P. knolli. The latter shows a mesoloph

in the upper molars that is absent in S. moyaeponsi. The upper

molars of P. knolli show a well-developed metaloph and a trans-

verse protoloph whereas in S. moyaeponsi the metaloph is absent

and the protoloph is obliquely directed. Contrary to the condi-

tion observed in the upper molars of P. knolli, the lophs of

S. moyaeponsi are very slender.

Paraphiomys simonsi Wood, 1968. The holotype and single

published specimen of this species (CGM 26908, Wood, 1968,

fig. 5F) is a badly damaged lower jaw with d4-m3 and the inci-

sor. It comes from the Oligocene portion of the Jebel el Qatrani

Formation at ‘Yale Quarry I’, Fayum Province, Egypt (Wood

1968). Interestingly, unpublished material of this species includes

a fragment of maxilla with P4-M2 (CGM 80–17) and two man-

dibular fragments, one of these with d4-m2 (CGM 40428) and

the other with m1-2 (CGM 40478). Paraphiomys simonsi is

one of the largest thryonomyids found to date. According to

Patterson and Wood (1982, p. 520), this species should be inclu-

ded in the genus Neosciuromys because of the high hypsodonty,

the simplification of the occlusal pattern and the large size of

the m2 and m3. However, according to recent cladistic analyses

TEXT -F IG . 2 . Paraphiomys knolli sp. nov. A, AS8-1100 (holotype), right m1-2. B, AS21-1101, left D4. C, AS8-1102, right M1-2. D,

AS8-1103, right M1-2. E, AS8-1104, left M3. F, AS8-1105, right M3. All figures are in occlusal view. Scale bar represents 1 mm.

L O P E Z A N T O N A N Z A S A N D S E N : N E W M I O C E N E R O D E N T 225

(Lopez-Antonanzas 2004; Lopez-Antonanzas et al. 2004), the

high hypsodonties of Neosciuromys africanus and P. simonsi are

homoplastic. The main differences between P. knolli and

P. simonsi are the hypsodonty and great size of the latter. The

lower molar of P. knolli shows a vestigial metalophulid II whereas

it is absent in P. simonsi. In P. simonsi, a small basal cingulum

lies along the anterior side of the protoconid, rising into a small

cusp, whereas P. knolli the anterolabial cingulum is lacking and

the anterior cusp is larger. Contrary to the upper molars of

P. knolli, those of P. simonsi have a three-lophed morphology.

Paraphiomys pigotti Andrews, 1914. This species was described

and illustrated by Andrews (1914, pp. 177–178) based on a frag-

mentary left mandible with p4-m2 from the Lower Miocene Bed

24 at Kachuku, Karungu area, Kenya. This taxon has been recor-

ded in the Lower Miocene of Rusinga (Lavocat 1973), Moruorot

(Madden 1972; Pickford 1981), and Gumba (Pickford 1981)

Kenya. It is also known in the Lower Miocene of Uganda at

Napak and Bukwa (Pickford 1981; Winkler and Downs 2002),

and it has been reported in the Middle Miocene in the Kenyan

localities of Fort Ternan (Shipman et al. 1981, p. 67) and Kiri-

mon (Pickford 1981; p. 89).

The morphology of the m1–2 of Paraphiomys pigotti (Andrews

1914, pl. 28, fig. 7; Lavocat 1973, pl. 27, fig. 9) is close to that of

the single m1-m2 recorded at As-Sarrar. However, the much

greater size of P. pigotti warrants its systematic distinction from

the As-Sarrar taxon. Furthermore, it shows the hypoflexus posteri-

orly directed whereas it is transverse in the m1–2 from As-Sarrar.

The latter also differs from P. pigotti in having the lingual end of

the posterolophid bent forwards, i.e. towards the lingual end of

the entoconid. Therefore, the metaflexus is more closed than that

of P. pigotti, in which this sinus is entirely open lingually. The m1-2

of P. pigotti (Andrews 1914, pl. 28, fig. 7; Lavocat 1973, pl. 27,

fig. 9), as well as that of P. knolli, show a short metalophulid II

that does not join the metaconid. Both taxa also have an isolated

and transversally elongated anteroconid. The upper molars of

P. pigotti mainly differ from those of P. knolli in their great size.

Lavocat (1973, p. 39) considered Phthinylla fracta Hopwood, 1929

to be a junior synonym of Paraphiomys pigotti, which was acknow-

ledged by Hendey (1978). However, the equally high hypsodonty,

and the discovery of Phthinylla-like maxillary teeth along with

Neosciuromys-like mandibles (P. Mein, pers. comm. 2003) suggest

that the former taxon is a junior synonym of the latter (Lopez-

Antonanzas 2004; Lopez-Antonanzas et al. 2004), as suspected by

Flynn and Winkler (1994) and Winkler (1994).

Only one of the two complete maxillae considered by Lavocat

(1973, p. 37) to be Paraphiomys pigotti corresponds to this taxon

(see Lavocat 1973, pl. 24, fig. 4); the other one (Phthinylla fract-

a-like) pertains to Neosciuromys africanus. The upper molars of

P. pigotti and those from As-Sarrar show a mesoloph, but this is

much less developed in the latter. It can even join the metacone

in P. pigotti. Furthermore, the mesoloph is posteriorly directed

in P. pigotti whereas it is nearly transverse in the specimens from

As-Sarrar.

Neosciuromys africanus Stromer, 1922. Neosciuromys africanus

has been described on the basis of material from the Lower Mio-

cene of Langental and Plant 4 borehole (south of Luderitz Bay,

Namibia). Stromer (1922, 1924, p. 263; 1926, pp. 135–136) des-

cribed and figured as N. africanus two mandibular fragments

with p4-m2 and a fragmentary left maxilla. Interestingly, the

latter specimen is actually a right mandibular fragment with

m2-m3 of the bathyergoidid Bathyergoides neotertiarius (Mein,

pers. comm. 2003). In contrast, the abnormally narrow M1 fig-

ured by Stromer (1926, pl. 42, fig. 24) as cf. Phiomys andrewsi

belongs instead to Neosciuromys africanus. There are obvious

morphological and dimensional differences between Neosciuro-

mys africanus and Paraphiomys knolli.

With regard to the lower molars, AS8-1100 is much smaller

than the m1–2 of N. africanus. In addition, Neosciuromys has

higher-crowned teeth and the pattern of the lower cheek teeth is

reduced to three transverse crests with no trace of metalophulid

II. In the specimen from As-Sarrar, the hypolophid is nearly

transverse whereas it is anteriorly directed in N. africanus. The

antero-posterior difference in width of AS8-1100 is much more

pronounced than in the lower molars of N. africanus. Finally,

the m1-2 from As-Sarrar shows a transverse hypoflexus whereas

it bends antero-posteriorly in N. africanus.

The single D4 from As-Sarrar shows a very short metaloph

whereas it is more developed in N. africanus. AS8-1101 is also

much smaller and less hypsodont than the D4 of N. africanus

and has the protoloph and metaloph anteriorly directed whereas

in N. africanus they are transverse. Contrary to the condition in

N. africanus, in the hypoflexus of AS8-1101 is nearly transverse.

The upper molars from As-Sarrar are very different from

those of N. africanus. As explained above, the maxilla described

by Hopwood (1929, p. 4, fig. 3) as Phthinylla fracta belongs to

N. africanus. The main differences between the As-Sarrar taxon

and N. africanus are the great size and hypsodonty of the latter.

Unfortunately no M3 of N. africanus has been described and

illustrated to date.

Apodecter stromeri Hopwood, 1929. The holotype specimen of

Apodecter stromeri (AMNH 22538) is a right mandibular ramus

with m1-m3, from an unknown Lower Miocene locality of the

Namib desert, south of Luderitz Bay (Hopwood 1929).

Lavocat (1973, p. 42), without justification, reallocated

Apodecter stromeri to the genus Paraphiomys. He noted (p. 44)

that the specimens of P. stromeri found at Rusinga (Kenya,

Lower Miocene) are smaller than those from Songhor (Kenya,

Lower Miocene) and lack the metalophulid II whereas the lower

molars from Songhor frequently show it. Somewhat unusually,

he thought that these differences were insufficient to justify a

specific distinction, but that they are nonetheless reliable indica-

tions of the existence of two subspecies of P. stromeri. Therefore,

he erected Paraphiomys stromeri stromeri to accommodate both

the material from Rusinga and the holotype specimen of Apodec-

ter stromeri, and Paraphiomys stromeri hopwoodi for the material

from Songhor.

We consider that the morphological differences between

these two ‘subspecies’ are sufficient to elevate them to species

rank (Lopez Antonanzas et al. 2004). However, there is no

reason to consider the smaller, more hypsodont and older

holotype of Apodecter stromeri and the remaining specimens of

Paraphiomys stromeri stromeri to be representatives of the same

species. In a very recent systematic study of the Thryonomyidae

226 P A L A E O N T O L O G Y , V O L U M E 4 8

(Lopez-Antonanzas 2004; Lopez-Antonanzas et al. 2004), the

genus Apodecter has been rehabilitated, the material from Ru-

singa has been considered a new species, Paraphiomys renelavo-

cati, and the material from Songhor has been designated as

P. hopwoodi.

The As-Sarrar material differs from Apodecter stromeri in hav-

ing a metalophulid II in the lower molars (absent in Apodecter)

and in the anterior part of the tooth being much wider than the

posterior one (it is only slightly wider in A. stromeri).

Unpublished material from the Namibian locality of Elisabeth

Felt (Mein in prep.) allows a comparison of the upper molars

of P. knolli with those of Apodecter. The basic morphology

of the upper dentition of both taxa is very close. However

the specimens of Apodecter are higher crowned than those from

As-Sarrar.

Paraphiomys hopwoodi Lavocat, 1973. The holotype of Paraph-

iomys hopwoodi (Lavocat 1973, p. 44) is a left mandibular frag-

ment (KNM SO 627) with d4-m2 and an erupting m3 from the

Lower Miocene of Songhor (Text-fig. 1). Although the lower

molars of P. hopwoodi are larger than AS-1008, they are very

similar in morphology. There are slight differences between the

upper molars of the two taxa: those of the As-Sarrar specimen

have a metaloph (absent in P. hopwoodi), a more marked antero-

posterior compression and a hypocone that is more labially

displaced.

Paraphiomys renelavocati Lopez-Antonanzas, Sen and Mein,

2004. The holotype of this species is a mandible with d4-m3

from the Early Miocene of Rusinga figured by Lavocat (1973,

pl. 26, fig. 9). It is housed in the National Museums of Kenya

(Nairobi) where it is available for study under reference number

KNM RU 2208. The dentition of this taxon is very similar in

size to specimens from As-Sarrar, but the morphology is differ-

ent enough in lacking a metalophulid II on the lower molars to

consider the latter to be distinct. Both taxa have a mesoloph on

the upper molars.

Kochalia geespei de Bruijn and Hussain, 1985. The holotype of

this species is a left d4 or p4 (p4 according to de Bruijn and

Hussain 1985, pl. 1, fig. 7a–b) from the middle part of the

Lower Manchar Formation of Pakistan (locality H-GSP 82.24).

The lower molars of Kochalia geespei mainly differ from the

As-Sarrar specimen in lacking any trace of metalophulid II and

in being more lophodont. The upper molars of the two taxa are

very similar. However, K. geespei usually lacks the metaloph

whereas in all of the As-Sarrar specimens it is complete. The

two taxa are comparable in size.

Kochalia sp. De Bruijn and Hussain (1985) also described a sec-

ond species of Kochalia on the basis of first and second right

lower molars from the base of the Nari (Early Miocene) Forma-

tion at the locality H-GSP 81.14A. According to them, this is a

different taxon from K. geespei because of its smaller size, lower

crowned cheek teeth, and the presence of metalophulid II on the

two lower molars. De Bruijn and Hussain (1985) pointed out

the close similarity between these teeth and the lower molar

from As-Sarrar and we concur with them in considering that the

similar size and morphology strongly advocate that all of these

specimens could be assigned to the same species.

Epiphiomys coryndoni Lavocat, 1973. The holotype of this

species (KNM RU 2253) is a left mandibular fragment with

m1-2 from the Lower Miocene of Rusinga, Kenya (Lavocat

1973, p. 46). According to Lavocat (1973), this species has

also been found at Songhor (Kenya) and Napak (Uganda)

(Text-fig. 1).

One of the main differences between Epiphiomys coryndoni

and AS8-1100 is the greater development of the metalophulid II

in the former. With regard to the upper molars, those of E. co-

ryndoni show a mesoloph that is much more developed than in

specimens from As-Sarrar. The molars of E. coryndoni are only

slightly smaller than those of Paraphiomys knolli.

Paraphiomys occidentalis Lavocat, 1961. This species has been

reported from two Middle Miocene sites in Morocco: Beni-

Mellal (Lavocat 1961, p. 45; Jaeger 1977, p. 121) and Azdal (Bena-

mmi et al. 1995) (Text-fig. 1). Little material of Paraphiomys

occidentalis is known to date. The type (UM Ben. Mel. 340) is a

right fragmentary maxilla with P4-M1 (Lavocat 1961, pl. 4,

fig. 1) from Beni-Mellal and the cotype (UM Ben. Mel. 667)

consists of a right m2. The Moroccan site of Azdal has yielded a

single right mandibular ramus (Benammi et al. 1995). The main

difference between P. occidentalis and the specimens from As-

Sarrar is in the size, the former being much larger than any

extinct thryonomyid, except Neosciuromys africanus, Paraphiomys

simonsi and P. pigotti. The lower molars of P. occidentalis and

those of P. knolli have a vestigial metalophulid II and an antero-

labial cuspid, which is larger in the latter. The posterolophid is

more curved in P. knolli than in P. occidentalis.

With regard to the upper molars, P. occidentalis shows an

unreduced mesoloph and lacks the metaloph, whereas those

from As-Sarrar show a mesoloph and a short but distinct

metaloph. The protoloph is transverse in the As-Sarrar speci-

mens whereas it curves posteriorly in P. occidentalis.

Paraphiomys shipmani Denys and Jaeger, 1992. This species

was described by Denys and Jaeger (1992, p. 77) based on five

mandibular rami from the Middle Miocene of Fort Ternan,

Kenya (Text-fig. 1). The holotype (KNM FT 3305) is a right

mandible with d4-m3. Unfortunately, no upper tooth of this

taxon has been found to date. The lower molars of Paraphiomys

shipmani differ from P. knolli in lacking a metalophulid II. Fur-

thermore, the anterior part of the tooth in the latter is wider

than the posterior part, whereas in P. shipmani they are nearly

equal in width. Contrary to the condition in AS8-1100, the

lower molars of P. shipmani show the hypolophid bending back-

wards. The posterolophid is more curved in P. knolli than in

P. shipmani, in which it is nearly transverse.

Paraulacodus indicus Hinton, 1933. Hinton (1933, pp. 621–

622) established the taxon Paraulacodus indicus on the basis of

three specimens (G.S.I. D281, D282, D283) from the Siwalik

‘series’ in Pakistan (then part of northern India). Only a short

diagnosis without illustrations was given, but Black (1972) pro-

vided a thorough systematic revision of the species, including

L O P E Z A N T O N A N Z A S A N D S E N : N E W M I O C E N E R O D E N T 227

original drawings of the three specimens available to Hinton

(1933).

The holotype of this species (GSI D283) is an incomplete

right maxilla with P4-M2 (Black 1972, fig. 2a–c) from an

unknown locality. An isolated upper molar of Paraulacodus aff.

Paraulacodus indicus from the Middle Miocene Chinji Formation

of Pakistan has been reported by Wessels et al. (1982, p. 358,

pl. 2, fig. 14). However, the great difference in size and slight

morphological differences between this tooth and equivalent

teeth of P. indicus suggest that it does not belong to this species

(de Bruijn and Hussain 1985; Lopez-Antonanzas 2004; Lopez-

Antonanzas et al. 2004). De Bruijn and Hussain (1985) designa-

ted it as Paraulacodus sp., an identification that we follow.

De Bruijn and Hussain (1984, 1985) reported specimens of

Paraulacodus cf. P. indicus from the Pakistani locality H-GSP

82.14 (top of the Lower Manchar Formation, Middle Miocene).

Later, the middle Chinji Formation of Pakistan yielded new

specimens of Paraulacodus indicus (Flynn and Winkler 1994).

There are some important differences between the lower

molars from As-Sarrar and those of Paraulacodus indicus. The

m1–2 of P. indicus are much larger than the unique lower molar

recovered from As-Sarrar and show three transverse crests and

no remnant of metalophulid II (whereas Paraphiomys knolli

shows this). In P. knolli , the anterior half of the crown is appre-

ciably wider than the posterior half, whereas in Paraulacodus

indicus the anterior half of the crown is only slightly wider than

the posterior. The upper molars from As-Sarrar are smaller than

those of P. indicus and show a mesoloph, which is absent in the

Pakistani form. Whereas all As-Sarrar specimens show a short

metaloph connecting the metacone with the posteroloph, the

upper molars of P. indicus lack it. The protoloph is transverse in

P. knolli, whereas it bends towards the labial end of the anterol-

oph in the upper teeth of P. indicus.

Paraulacodus johanesi Jaeger, Michaux and Sabatier, 1980. The

holotype of this species (NMAA CHO 1) was described by Jaeger

et al. (1980, p. 367). It is an incomplete maxilla with P4-M2

(Jaeger et al. 1980, fig. 1a) from the Upper Miocene of Chorora,

Ethiopia (Jaeger et al. 1980). Additional material was found later

at the same locality (Geraads 1998, 2001, p. 104) and in the

Upper Miocene of northern Kenya in the lower member of the

Nawata Formation (Winkler 2003) and possibly in the Namu-

rungule Formation (Kawamura and Nakaya 1984, 1987). The

existence of a metalophulid II in AS8-1100 and the much larger

size of Paraulacodus johanesi suggest that there are no close rela-

tionships between them. In addition, contrary to the condition

in P. johanesi, the unique lower molar from As-Sarrar shows a

distinct, isolated anteroexternal cusp. Furthermore, its hypoflex-

us is transverse whereas in P. johanesi it is obliquely directed.

With regard to the upper molars, those of the As-Sarrar form

show a mesoloph, which is absent from those of P. johanesi. In

addition, the posterior half of the crown is relatively wider in

Paraphiomys knolli than in P. johanesi.

Paraphiomys afarensis Geraads, 2001. Geraads (1998, p. 210)

based his new species Paraphiomys chororensis on two upper

molars, two fragmentary upper molars, five fragmentary mandi-

bles, some isolated lower molars and several incisors, all from

the early Late Miocene Ethiopian locality of Chorora (Geraads

et al. 2002). The holotype is an upper molar (unnumbered)

housed in the National Museum of Addis Ababa (Geraads 1998,

pl. 2, fig. 6) and corresponds to the so-called Paraphiomys sp. 2

of Jaeger et al. (1980, p. 371, fig. 3a–b). New material collected

at Chorora has revealed that the holotype of P. chororensis is

actually a P4 of Paraulacodus johanesi. Geraads (2001, p. 103)

coined the name Paraphiomys afarensis to accommodate the

remaining specimens of the hypodigm of P. chororensis. This

species has been recently recorded from the Late Miocene of the

lower member of the Nawata Formation, Lothagam, Kenya

(Winkler 2003).

The morphology of the lower teeth of Paraphiomys afarensis

is very different from that of P. knolli. The three-lophed lower

molars of P. afarensis clearly differ from the four-lophed AS8-

1100. According to Geraads (1998, p. 210), P. afarensis retains

the mesoloph in the upper molars, which is also obvious in the

upper molars from As-Sarrar. With respect to size, the teeth of

P. afarensis are much larger than those of P. knolli.

Paraphiomys roessneri Mein, Pickford and Senut, 2000. The hol-

otype of this species is an upper left molar (GSW Ari 2) from

the Upper Miocene Harasib deposits of Namibia. Paraphiomys

roessneri is distinguished from the As-Sarrar specimens by the

absence of metalophulid II and in lacking a prominent labial

anteroconid on the lower molars. With regard to the upper

molars, one of the more important differences between the two

taxa is the absence of a mesoloph in the teeth of P. roessneri.

Paraphiomys australis Mein, Pickford and Senut, 2000. This

species was erected on the basis of 317 isolated teeth and two

maxillae from the Upper Miocene Harasib deposits of Namibia.

The holotype of this species (GSW Ari 13) is a right p4 (Mein

et al. 2000, fig. 1.13). In addition to the larger size of the teeth of

Paraphiomys australis in comparison with those from As-Sarrar,

this species differs notably in the absence of metalophulid II and

in lacking the anterior cusp on the lower molars. With regard to

the upper molars, one of the differences between the two taxa is

the absence of mesoloph in P. australis.

Paraphiomys orangeus Mein and Pickford, 2003. This species

has been reported from the Early Miocene (17Æ5–17 Ma) Orange

River deposit of Arrisdrift, Namibia (Mein and Pickford 2003,

p. 153). The holotype, AD 408¢97, is a left mandible with i, p4-m3

(Mein and Pickford 2003, fig. 7). This species differs from Para-

phiomys knolli in lacking metalophulid II on the lower molars.

SYSTEMATICS AND PHYLOGENY

McKenna and Bell (1997, pp. 194–195) included many

genera in the Thryonomyidae. However, conflicting opin-

ions have been expressed on the familial attribution of

Paraphiomys, Neosciuromys, Apodecter, Gaudeamus, Epi-

phiomys and Sacaresia. Thus, Paraphiomys is from time to

time seen as a taxon of the extinct family Phiomyidae

(Wood 1968; de Bruijn 1986; Denys and Jaeger 1992;

Geraads 1998). The same is true for Neosciuromys (Wood

228 P A L A E O N T O L O G Y , V O L U M E 4 8

1968), Apodecter (Wood 1968), Gaudeamus (Wood 1968)

and Epiphiomys (de Bruijn 1986). In addition, Sacaresia

has been considered as a taxon close to, but outside,

the thryonomyids by Flynn and Winkler (1994). Our

knowledge of the systematics and phylogeny of the

thryonomyids has recently been significantly improved

(Lopez-Antonanzas 2004; Lopez-Antonanzas et al. 2004).

With the purpose of finding the phylogenetic position

of Paraphiomys knolli within the family Thryonomyidae, a

cladistic analysis has been conducted. In addition to

P. knolli, the taxa involved in this analysis are the rodent

species commonly referred to as thryonomyid representa-

tives whose the validity has been established by Lopez-

Antonanzas (2004) and Lopez-Antonanzas et al. (2004),

namely Apodecter stromeri, Epiphiomys coryndoni, Gaude-

amus aegyptius, Kochalia geespei, Neosciuromys africanus,

Paraphiomys afarensis, P. australis, P. hopwoodi, P. occi-

dentalis, P. orangeus, P. pigotti, P. renelavocati, P. roessneri,

P. shipmani, P. simonsi, Paraulacodus indicus, P. johanesi,

Sacaresia moyaeponsi, Thryonomys gregorianus and

T. swinderianus.

The addition of the non-Thryonomyidae Metaphiomys

schaubi permits us to test the monophyly of the group

constituted by the species listed above. The outgroup

taxon Phiomys andrewsi (Phiomyidae) gives a direction to

the character transformation and an order of reading of

the successive branchings of the tree. Previous cladistic

analyses (Marivaux 2000; Marivaux et al. 2002) showed

that Phiomys is external to a clade comprising Paraphio-

mys and Paraulacodus. Interestingly, Phiomys does not

show a morphological gap with the taxa of the ingroup

and it was selected as outgroup in the phylogenetic analy-

sis of thryonomyid relationships presented by Winkler

(1992) and Flynn and Winkler (1994).

Twenty-two phylogenetically informative dental charac-

ters are coded. Fifteen characters are binary whereas seven

are multistate. Among the latter, characters 5, 12–14, 19

and 22 have two derived states and character 17 has three.

Owing to the lack of a priori information, the weight of

all characters is identical (1), and there is no rationale to

order the multistate characters; hence, all the possible

transformations of one state into another will cost one

step.

The software of phylogenetic reconstruction by parsi-

mony analysis used is PAUP version 3.1.1 (Swofford

1993). The number of terminal taxa and characters has

precluded the exact tree-building method, so a heuristic

search with stepwise addition (‘closest’ option) was per-

formed.

Four most parsimonious trees (80 steps long) have resul-

ted. Their characteristics are CI ¼ 0Æ375, RI ¼ 0Æ580. The

strict consensus tree is identical to that of the majority-rule

with two irresolutions (nodes 43 and 27). The semi-strict

consensus tree (Text-fig. 3) supports 20 internal nodes: it

TEXT -F IG . 3 . Semi-strict consensus tree generated by the cladistic analysis of the Thryonomyidae, showing phylogenetic position of

Paraphiomys knolli sp. nov. Dashed line indicates the node that collapses in the strict consensus tree. Length ¼ 80 steps, Consistency

Index ¼ 0Æ375, Retention Index ¼ 0Æ580.

L O P E Z A N T O N A N Z A S A N D S E N : N E W M I O C E N E R O D E N T 229

only shows basal irresolution with regard to the respective

placement of Metaphiomys schaubi, Sacaresia moyaeponsi,

and the remaining taxa of the ingroup.

One of the most interesting results of the cladistic ana-

lysis presented herein is the evidence of the polyphyletic

nature of the genus Paraphiomys as currently used. In

fact, only a handful of thryonomyid species are narrowly

related to P. pigotti, the type species of the genus, and

therefore could legitimately continue to be attributed to

it. These are P. knolli, P. hopwoodi, P. occidentalis and

P. renelavocati.

The monophyly of the two species of Paraulacodus

(node 26) is only present in 50 per cent of the most parsi-

monius trees. This monophyly is supported by the results

of the phylogenetic analysis of Winkler (1992) and Flynn

and Winkler (1994).

Based on extrinsic (age) considerations the branching

of Gaudeamus aegyptius in the crown group is unexpec-

ted. Lavocat (1973, p. 46) considered Gaudeamus to be

more progressive than Thryonomys. By contrast, Wood

(1968, fig. 17) considered Gaudeamus aegyptius to be a

direct ancestor of the Recent Thryonomys. Incidentally, de

Bruijn and Whybrow (1994) showed a thryonomyid m3

from Abu Dhabi (United Arab Emirates) that morphologi-

cally recalls both Gaudeamus and Thryonomys, and is

intermediate from both size and age (Late Miocene) points

of view. Therefore it turns out that Gaudeamus aegyptius

should be reallocated to the genus Thryonomys (a taxo-

nomic alternative compatible with the present cladogram

topology would be to restore the genus Choeromys for

Thryonomys gregorianus, as initiated by Thomas 1992).

Nevertheless, this part of the cladogram implies that there

is a missing lineage duration of about 30 myr. The posi-

tion of Gaudeamus aegyptius within species of Thryonomys

could be due to convergent evolution towards the same

kind of simplified cheek teeth. In fact, the incisors and

jaws of Gaudeamus and Thryonomys look nothing like

each other and the entire body of the former was about

the size of the head of a Thryonomys. Formally synony-

mizing Gaudeamus with Thryonomys would definitely

place too much confidence in the proposed phylogeny.

CONCLUSION

Winkler (1992, p. 244) suggested that the thryonomyid

rodent from As-Sarrar represents ‘a new species or poss-

ibly a new genus’. Whereas the first possibility has been

confirmed by our study, the existence of a new genus has

been dismissed. In fact, Paraphiomys knolli cannot be dis-

tinguished, in the present state of knowledge, from the

taxon referred to as ‘Kochalia sp.’ by de Bruijn and Huss-

ain (1985). Therefore, we consider provisionally that it is

the same as that from the Early Miocene Nari Formation

in Pakistan. However, it cannot be considered as conge-

neric with Kochalia geespei. Whereas the latter is quite

plesiomorphic (sister-group of the unnamed clade com-

prising ‘Paraphiomys’ simonsi and all other more derived

thryonomyids), P. knolli branches as the sister-species of

P. pigotti, the type species of the genus. Paraphiomys hop-

woodi constitutes the sister-group of (P. pigotti, P. knolli).

As these three species are morphologically fairly close,

there is little doubt that the latter must be considered to

belong to the genus Paraphiomys.

As indicated by de Bruijn and Hussain (1985, p. 162),

the presence of the same species of rodent in the Arabian

Peninsula and the Indian subcontinent provides evidence

for the absence of barriers precluding faunal exchange

between these two distant zones during the first half of the

Miocene. This is supported by the geographical distribution

of the more abundant species of Sayimys during the Early

and Middle Miocene. Paraphiomys represents an important

clade of thryonomyid rodents of this time. Although still

lacking in the Miocene fossil record of some areas such as

Turkey, representatives of this genus were without doubt

widely distributed from western and northern Africa to the

Indian subcontinent before vanishing apparently without

having given rise to any descendants.

Acknowledgements. D. Batten (University of Wales, Aberyst-

wyth), L. Flynn (Harvard University, Cambridge, USA), P. Hol-

royd (University of California, Berkeley), F. Knoll (Staatliches

Museum fur Naturkunde, Stuttgart) and P. Mein (Universite

Claude Bernard-Lyon 1, Villeurbanne) improved various aspects of

this paper through critical reading and ⁄ or discussion. A. Currant

(The Natural History Museum, London) kindly made available

for examination the holotype and paratypes of Paraulacodus

indicus, which are currently housed in his institution. The rodent

remains from Saudi Arabia were collected during field work

organized by H. Thomas (College de France, Paris) with permis-

sion of the Department of Antiquities and Museums in Riyadh.

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APPENDIX

List of characters used in this analysis.

Fifteen characters are binary whereas seven are multistate. The

plesiomorphic state (0) was obtained by outgroup comparison

with Phiomys andrewsi.

dp4

1. (0) replaced; (1) not replaced.

2. Anterior cingulum: (0) present; (1) weak or absent.

3. Metalophulid II: (0) present; (1) absent.

4. Metalophulid II: (0) transverse; (1) oblique.

5. Hypolophid: (0) posteriorly oblique; (1) transverse; (2) anteri-

orly oblique.

m1)2

6. Anterolabial cuspid: (0) absent; (1) present.

7. Anterolabial cuspid: (0) isolated; (1) partially submerged in

crest.

8. Metalophulid II: (0) present; (1) absent.

9. Mesoflexid: (0) not anteriorly directed; (1) anteriorly directed.

10. Mesoflexid: (0) equal or longer than the metaflexid; (1)

shorter than the metaflexid.

11. Anterolabial cingulum: (0) present; (1) weak or absent.

12. Hypolophid: (0) posteriorly oblique; (1) transverse; (2) ante-

riorly oblique.

13. Length: (0) m1 < m2; (1) m1 ¼ m2; (2) m1 > m2.

14. Width: (0) m1 < m2; (1) m2 ¼ m1; (2). m1 > m2.

m3

15. Length: (0) m3 > m2; (1) m3 < m2.

16. Anterolabial cingulum: (0) present; (1) weak or absent.

I

17. (0) ungrooved upper incisor; (1) one groove, (2) two

grooves, (3) three grooves.

M1)2

18. Mesoloph: (0) present; (1) absent.

19. Length: (0) M1 < M2; (1) M1 ¼ M2; (2) M1 > M2.

20. Metaloph: (0) present; (1) absent.

21. Protoloph: (0) transverse; (1) oblique.

22. Hypsodonty: (0) low; (1) semihypsodonty; (2) high.

Matrix of character codings used in the analysis of

relationships of all species of Thryonomyidae

Characters are listed above. Phiomys andrewsi is used as out-

group. Character codings: 0, postulated plesiomorphic condition

of character; 1, 2 and 3, postulated derived conditions of charac-

ter; –, uncodable character; ?, character state uncertain.

232 P A L A E O N T O L O G Y , V O L U M E 4 8

Taxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22

Phiomys andrewsi 0 0 0 0 0 0 - 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Metaphiomys schaubi 0 0 0 1 2 0 - 0 1 0 0 2 0 0 0 0 0 0 0 0 0 0

Gaudeamus aegyptius 0 1 0 0 0 0 - 1 1 1 1 0 0 0 1 1 ? 1 0 1 1 1

Sacaresia moyaeponsi 0 0 0 1 0 0 - 0 0 0 0 0 0 0 1 0 1 1 0 1 1 0

Paraphiomys simonsi ? 1 1 - 2 1 0 1 1 0 1 0 0 0 0 0 0 0 0 1 0 2

Paraphiomys pigotti 1 1 1 - 2 1 0 0 0 1 1 2 0 0 0 1 0 0 1 0 0 1

Neosciuromys africanus 1 1 1 - 2 1 1 1 0 0 1 2 0 0 0 1 ? 0 0 0 1 2

Paraphiomys knolli sp. nov. ? ? ? ? ? 1 0 0 0 0 1 2 ? ? ? 1 ? 0 ? 0 0 0

Apodecter stromeri 1 1 1 - 2 1 1 1 0 0 0 1 2 2 1 1 0 0 0 0 0 1

Paraphiomys orangeus 1 0 1 - 1 1 1 1 1 0 1 1 0 0 1 1 0 0 0 0 0 1

Paraphiomys hopwoodi 1 1 1 - 2 1 0 0 0 0 1 2 1 0 1 1 0 0 ? 1 1 1

Paraphiomys renelavocati 1 1 1 - 2 1 1 1 0 0 0 1 0 1 1 1 0 0 1 1 1 1

Epiphiomys coryndoni ? 0 0 1 2 1 0 0 0 0 0 1 2 0 1 1 ? 0 2 0 0 1

Paraphiomys occidentalis 1 1 1 - 2 1 0 1 0 0 1 1 0 0 0 1 ? 0 2 1 1 1

Paraphiomys shipmani 1 1 1 - 0 1 1 1 1 0 1 0 2 0 1 ? ? ? ? ? ? 0

Kochalia geespei 0 1 1 - 1 1 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 1

Paraulacodus indicus 1 1 1 - 1 1 1 1 0 0 1 1 2 0 1 1 2 1 0 0 1 1

Paraulacodus johanesi 1 1 1 - 1 1 1 1 0 0 1 1 0 0 1 1 2 1 0 0 1 1

Paraphiomys afarensis 1 1 1 - 2 1 1 1 0 0 1 0 0 0 0 1 0 0 ? 0 1 1

Paraphiomys roessneri 1 0 1 - 1 0 - 1 0 0 0 0 0 2 1 1 0 1 2 0 0 1

Paraphiomys australis 1 0 1 - 2 0 - 1 0 0 0 0 2 2 1 0 ? 1 0 0 1 1

Thryonomys swinderianus 1 1 0 1 1 0 - 1 0 0 1 1 0 0 1 1 3 1 0 1 1 1

Thryonomys gregorianus 1 1 0 1 1 0 - 1 0 0 1 0 0 0 1 1 3 1 0 1 1 1

L O P E Z A N T O N A N Z A S A N D S E N : N E W M I O C E N E R O D E N T 233