Fossils and palaeontological distributions of Macaranga and Mallotus (Euphorbiaceae)

Abstract

The correct identification of described plant fossils from the sister genera Macaranga and Mallotus (Euphorbiaceae) needs to be confirmed in order to correctly date their phylogeny and map their palaeontological distributions. Previous identifications of fossil specimens often appear to be unreliable since the distinctive features for identification such as leaf details and stamen thecae are usually not preserved in the fossils. Fossil pollen, seeds, wood and leaf (imprints) are so general in appearance that they are only typical either for both genera and their sister group Hancea (pollen), the family Euphorbiaceae (seeds) or even a large group of families (wood, leaf venation). Out of 54 fossils only four could be reliably identified as Macaranga and Mallotus, which is sufficient for dating the phylogeny, but insufficient to provide an insight into palaeontological distributions. All discussed fossils are mapped in order to assess the possibility of ancestral distributions outside the presently known distribution of Macaranga and Mallotus, which otherwise is impossible due to the limitations of the current historical biogeographic techniques that are based on present day distributions. A short overview of best practice to evaluate fossils and their identification is presented.

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Fossils and palaeontological distributions of Macaranga and Mallotus (Euphorbiaceae)
M. Nucete, J.H.A. van Konijnenburg-van Cittert, P.C. van Welzen ⁎
Netherlands Biodiversity Centre Naturalis, section National Herbarium of the Netherlands, Leiden University, P.O. Box 9514, 2300 RA Leiden, The Netherlands
abstractarticle info
Article history:
Received 26 March 2012
Received in revised form 29 June 2012
Accepted 7 July 2012
Available online 24 July 2012
Keywords:
Euphorbiaceae
Macaranga
Macrofossils
Mallotus
Microfossils
Palaeontological distributions
The correct identification of described plant fossils from the sister genera Macaranga and Mallotus
(Euphorbiaceae) needs to be confirmed in order to correctly date their phylogeny and map their
palaeontological distributions. Previous identifications of fossil specimens often appear to be unreliable
since the distinctive features for identification such as leaf details and stamen thecae are usually not pre-
served in the fossils. Fossil pollen, seeds, wood and leaf (imprints) are so general in appearance that they are
only typical either for both genera and their sister group Hancea (pollen), the family Euphorbiaceae (seeds)
or even a large group of families (wood, leaf venation). Out of 54 fossils only four could be reliably identified
as Macaranga and Mallotus, which is sufficient for dating the phylogeny, but insufficient to provide an insight
into palaeontological distributions. All discussed fossils are mapped in order to assess the possibility of ancestral
distributions outside the presently known distribution of Macaranga and Mallotus, which otherwise is impossi-
ble due to the limitations of the current historical biogeographic techniques that are based on present day
distributions. A short overview of best practice to evaluate fossils and their identification is presented.
© 2012 Elsevier B.V. All rights reserved.
1. Introduction
Macaranga Thou. and Mallotus Lour. are two large genera (240 and
110 spp., respectively), which were recently revised, Macaranga by
Whitmore (2008), and Mallotus by a group of researchers
(Bollendorff et al., 2000; Slik and van Welzen, 2001; Sierra and Van
Welzen, 2005; Sierra et al., 2005, 2007a; Van Welzen and Sierra,
2006; Van Welzen et al., 2006, 2010). A core phylogeny of both gen-
era was recently published (Kulju et al., 2007a; Sierra et al., 2010).
The two genera appear to be sister groups when part of Mallotus is ex-
cluded and combined with the genus Cordemoya Baill. (Sierra et al.,
2006) as the genus Hancea Seem. (Sierra et al., 2007b), and when
the genera Neotrewia Pax & K.Hoffm., Octospermum Airy Shaw, and
Trewia L. are united with Mallotus (Kulju et al., 2007b).
Macaranga and Mallotus are morphologically very similar; the
only real difference is the number of thecae in the stamens (3 or 4
thecae in Macaranga,2inMallotus). Usually, Mallotus has stellate
hairs or stellately grouped simple hairs, while the hairs in Macaranga
are generally simple only. Even far less absolute differences can be
found in the inflorescences (more paniculate in Macaranga, more
spiciform in Mallotus) and the staminate flowers (usually fewer sta-
mens in Macaranga). Another partial difference between Macaranga
and Mallotus is in the leaf arrangement; Macaranga leaves are always
alternate, whereas Mallotus also has species with opposite leaves.
Typical for both genera is the presence of small glandular
scale-like hairs (generally on the lower leaf surfaces, inflorescences,
flowers and fruits; see figures A10–A12 in Sierra et al., 2010), domatia
(small structures, e.g., dots of hair, on the lower surface of the leaf,
usually at the junction of the midrib and the secondary nerves) and
extrafloral nectaries on the upper surface near the base and/or top
of the leaves (not in the central part of the leaf blade). The species
are usually dioecious and the flowers are of a very simple construc-
tion, a few sepals and stamens in the staminate flowers, and sepals
and a usually 3-locular gynoecium in the pistillate flowers. The fruits
are often spiny with soft spines. Phylogenies of both genera were
published by Kulju et al. (2007a) and Sierra et al. (2010) and for the
myrmecophytic Macaranga species (species living in obligate symbio-
sis with ants, which house in the stems and protect the plant) by
Blattner et al. (2001) and Bänfer et al. (2004).
Both genera also occur in similar habitats and they seemingly
responded in the same morphological way to changes in ecological
niches (small and narrow leaves in primary forest surroundings
changing to large and broad leaves in pioneer habitats; Slik et al.,
2003). The two genera also show a more or less similar distribution.
They are present in the central, tropical part of Africa, Madagascar,
the Mascarene Islands, south and southeast Asia from India and Sri
Lanka eastwards, whereby Mallotus reaches the main Japanese
Islands, while Macaranga has its northern range already in southern
China. The two genera are further present throughout the Malay
Archipelago up to the tropical forest region in eastern Australia and
the island chains in the West Pacific. They are absent from New
Zealand (Fig. 1). In all areas where both genera occur Macaranga gen-
erally has more species than Mallotus.
Palaeogeography, Palaeoclimatology, Palaeoecology 353–355 (2012) 104–115
⁎Corresponding author. Tel.: +31 715273571; fax: +31 715273511.
E-mail address: Welzen@nhn.leidenuniv.nl (P.C. van Welzen).
0031-0182/$ –see front matter © 2012 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2012.07.016
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It is intriguing that quite a number of fossils have been identified as
Macaranga and Mallotus, most within the present day distributions, but
also a number far outside the contemporary distribution (S. America,
N. America, Europe and Russia). The latter are questionable, because
Macaranga and Mallotus are thought to have mainly speciated relative-
ly recently (Kulju et al., 2007b; Sierra et al., 2010) after the split up of
Gondwana.
Fossils are generally used to date phylogenies, but only reliable
fossils can be considered as calibration points. Thus, the degree of
certainty with which the fossils are identified as Macaranga and/or
Mallotus will be evaluated.
We have the following three research questions:
−Which fossil Macaranga and Mallotus plant parts have been
described and how reliable are these identifications?
−Which fossils can be used to date a phylogeny?
−Was the distribution of Macaranga and Mallotus wider in former
geological times than presently?
2. Materials and methods
Literature of fossils described as Macaranga and Mallotus was
extensively collected via the second author (palaeobotanist) and her
network of experts, internet, and literature references in articles
pertaining to Macaranga and Mallotus fossils. In total 54 fossils were
listed in 44 articles. The literature was carefully read and analyzed
(Table 1). Most fossils appeared to be only small conserved parts of
plants; either leaf fragments, pollen, pieces of wood, or seeds. These
will be discussed separately. The following additional experts were
consulted: Dr. Max M.J. van Balgooy and Dr. Raymond W.J.M. van
der Ham (Herbarium Leiden University; leaves and pollen, respec-
tively), Dr. Wolfgang Stuppy (Royal Botanic Gardens, Kew; seeds)
and Dr. Elizabeth A. Wheeler (Dep. Wood and Paper Science, North
Carolina State University; wood). If the plates in the articles were
insufficient, additional high-resolution pictures were requested from
the authors when possible.
The following criteria were applied to evaluate the identifications
of the fossils:
−Diagnostic characters visible (e.g., flowers, fruits, extrafloral
nectaries and glandular scale-like hairs on leaves): then identifica-
tion is probably correct.
−No diagnostic characters visible, only general characters (e.g.,
venation pattern), but fossil present within present day distribu-
tion: then the fossils are classified as ‘unidentifiable within the
present day distribution’(UIPDD).
−No typical characters visible and fossil outside present day distribu-
tion: then the fossils are classified as ‘unidentifiable outside the
present day distribution’(UOPDD).
The palaeontological maps used to depict the sites where the
fossils were found and possibly where they originated, were created
by Dr. Ronald Blakey (Blakey, 2011)
Morphological similarities with modern species of Macaranga and
Mallotus present in Africa and for New Zealand from surrounding
areas (Australia, Pacific Island chains) were used to link fossils with
species or groups of species to which the fossils may be related and
which can then be used in the future to date the phylogeny.
3. Results
3.1. Leaves
Literature on 41 different fossil remains of leaves is evaluated in
this paper. The ages of the fossil leaves range from the Maastrichtian
(70 Ma) to the Pleistocene (1.8 Ma and younger). The results of the
evaluation of the fossils can be found in Table 1.
Most fossils show parts of leaves with only an imprint of a general
venation pattern, pinnate or basally 3-veined, generally secondary
veins looping and connecting near an entire or serrate margin. These
were too inadequate to identify as Macaranga and Mallotus (much
less identify them as either Macaranga or Mallotus), not only due to
the lack of typical characters (glandular hairs, domatia, and extra floral
Fig. 1. Present day distribution of Macaranga (black line) and Mallotus (white line) and locations of fossils attributed to these genera; Δ=Macaranga fossils; ○=Mallotus fossils;
□=Macaranga or Mallotus fossil; Green= identified fossil; Yellow =Unidentifiable within the Present Day Distribution (UIPDD) Red = Unidentifiable Outside the Present Day
Distribution (UOPDD).
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Table 1
Results for the evaluated literature on Macaranga and/or Mallotus fossils. Details in the column Evalution means anatomical details; only name mentioned means that no descriptions, drawings or photos were present.
Author(s) Year Fossil Species Location Time Evaluation Verdict
Ablaev, A.G., Vassiljev, I.V. 1994 Leaves Mallotus rjufflei Ablaev et Ig. Vassiljev E. Russia Oligocene Complete leaf, only venation, no details UOPDD
Akhmetiev, M.A., Vikulin, S.V. 1995 Leaves Macaranga peltata Müll.Arg. E. Russia Oligocene Complete leaf, only venation, no details UOPDD
Antal, J.S., Awasthi, N. 1993 Leaves Mallotus kalimpongensis Anatal & Awasthi Bengal, India Miocene–Pliocene Complete leaf, only venation, no details UIPDD
Antal, J.S., Awasthi, N. 1993 Leaves Macaranga siwalika Anatal & Awasthi Bengal, India Miocene–Pliocene Fragment, bad material and photos, no details UIPDD
Bande,M.B., Srivastava, G.P. 1990 Leaves Mallotus philippensis (Lam.) Müll.Arg. Bihar, India Miocene–
Pleistocene
Leaf fragment, only venation, no details UIPDD
Bonnefille, R. et al. 1987 Pollen Macaranga kilimandscharica Pax Hadar, Ethiopia Pliocene Only species mentioned. UIPDD
Bonnefille, R., Riollet, G. 1988 Pollen Macaranga type Burundi, Africa Holocene Only name mentioned. UIPDD
Cavagnetto, C., Anadon, P. 1996 Pollen Mallotus type NE Spain Eocene–Oligocene Only name mentioned UOPDD
Cavagnetto, C., Anadon, P. 1995 Pollen Mallotus type NE Spain Eocene Only name mentioned UOPDD
Currano, E.D. et al. 2011 Leaves Macaranga type NW Ethiopia, Africa Oligocene Leaf fragment, only venation, no details UIPDD
Fosberg, F.R., Corwin, G. 1958 Leaves Macaranga thompsonii Merr. Pagan, Mariana Islands Miocene–
Pleistocene
Only description, no anatomical details UOPDD
Garcia Massini, J.L. et al. 2010 Leaves Macaranga sp. Ethiopia, Africa Oligocene Complete leaf with anatomical details Correct?
Gruas-Carvagnetto, C.,
Kohler, E.
1992 Pollen Mallotus type N. France Eocene Description and not too clear photos. UOPDD
Guleria, J.S., Srivastava, R. 2001 Wood Mallotoxylon keriense Lakhanpal & Dayal W. India Maastrichtian Description and reasonable photos UIPDD
Horiuchi, J., Takimoto, H. 2001 Leaves Mallotus sp. Kanto, Japan Miocene Description, leaf fragment, only venation, no details UIPDD
Hu, H.H., Chaney, R.W. 1940 Leaves Mallotus populifolia Hu & Chaney Shantung, China Miocene Description, leaves, only venation, no details UIPDD
Khan, M.A. et al. 2011 Leaves Macaranga cf. denticulate (Blume) Müll.Arg. Arunachal, Pradesh, India Pliocene–Pleistocene Leaf fragment, only venation, no details UIPDD
Kramer, K. 1974 Wood Mallotoxylon keriense Lakhanpal & Dayal Central India Miocene Description, drawings without details UIPDD
Lakhanpal, R.N., Dayal, R. 1963 Wood Mallotoxylon keriense Lakhanpal & Dayal Central India Eocene Description, photos of not too well preserved wood UIPDD
Lee, D.E. et al. 2010 Leaves Malloranga fouldenensis Lee, Bannister, Raine & Coran Otago, New Zealand Miocene Full leaves, full anatomical details Correct?
Lee, D.E. et al. 2010 Fruit Malloranga fouldenensis Lee, Bannister, Raine & Coran Otago, New Zealand Miocene Damaged fruits, photos and description Correct?
Lee, D.E. et al. 2010 Pollen Nyssapollenites endobalteus (McIntyre) Kemp and Harris Otago, New Zealand Miocene SEM and LM photos, description Correct?
MacGinitie, H.D. 1941 Leaves Mallotus riparius MacGinitie Seirra Neveda, U.S.A. Eocene Full leaves, general venation, no details UIPDD
Mai, D.H. 2001 Seeds Mallotus tuberculatus Mai E. Germany Miocene Not too detailed photos, description UOPDD
Maley, J., Brenac, P. 1997 Pollen Macaranga type W. Cameroon Holocene Name mentioned UIPDD
Maley, J., Brenac, P. 1997 Pollen Mallotus type W. Cameroon Holocene Name mentioned UIPDD
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Author(s)
Year Fossil Species Location Time Evaluation Verdict
Mathur, A.K. 1978 Leaves Mallotus sp. Jawalamukhi, India Miocene Leaf fragment, general venation, no details, bad photo UIPDD
Martin, H.A. 1974 Pollen Macaranga–Mallotus New South Wales, Australia Eocene–Pliocene Name mentioned UIPDD
Mildenhall, D.C. 1980 Pollen Macaranga/Mallotus New Zealand Oligocene–Pliocene Name mentioned UOPDD
Pathak, N.R. 1969 Leaves Mallotus philippensis (Lam.) Müll.Arg. Darjeeling, India Pliocene-Pleistocene Description, bad quality leaf fragments and photos UIPDD
Pole, M. 1996 Leaves Macaranga/Mallotus Otago, New Zealand Miocene Description, drawing of bad leaf fragment, no details Correct?
Prakash, U., Tripathi, P.P. 1975 Wood Mallotoxylon assamicim Prakash & Tripathi E. India Miocene Description, one non-informative drawing, several
low quality photos of reasonable quality fossil
UIPDD
Prasad, M 1994 Leaves Mallotus venkatachalai Prasad Uttar Pradesh, India Miocene Leaf fragments, only venation, no details, bad photos UIPDD
Prasad, M., Awasthi, N. 1996 Leaves Mallotus venkatachalai Prasad W. Nepal Miocene Leaf fragment, only venation, no details, bad photo UIPDD
Prasad, M., Pandey, S.M. 2008 Leaves Mallotus kalimpongensis Anatal & Awasthi Himalaya, Nepal Miocene–Pliocene Description, leaves, only venation, no details. UIPDD
Puri, G. S. 1947 Leaves Mallotus philippensis (Lam.) Müll.Arg. Kashmir, India Pleistocene Leaf fragments, venation, no details, bad photos UIPDD
Rásky, K. 1965 Leaves Macarangaephyllum palaeomonandrum Rásky N. Hungry Eocene Leaf fragments, venation, no details, bad photos UOPDD
Rásky, K. 1965 Flower Antholithus macarangaeformis Rásky N. Hungry Eocene Small bad quality photo, no details, not interpretable UOPDD
Rásky, K. 1965 Leaves Mallophyllum palaeomiquelianum Rásky N. Hungry Eocene Leaf fragments, venation, no details, bad photos UOPDD
Roy, S.K., Ghosh, P.K. 1982 Wood Mallotoxylon cleidioniodes Rásky W. India Miocene Description, low quality fossil, bad photos UIPDD
Sanborn, E.I. 1937 Leaves Mallotus comstocki Sanborn Oregon, U.S.A. Eocene Description, leaf fragments, venation, no details UOPDD
Sanborn, E.I. 1937 Leaves Mallotus oregonensis Sanborn Oregon, U.S.A. Eocene Description, leaf fragments, venation, no details UOPDD
Tanai, T. 1970 Leaves Mallotus hokkaidoensis Tanai Hokkaido, Japan Oligocene Description, leaf, venation only, no details UIPDD
Tanai, T. 1989 Leaves Mallotus hokkaidoensis Tanai Hokkaido, Japan Eocene–Miocene Idem as former, veinlet drawings added, anatomical details
described
Correct?
Tanai, T. 1990 Leaves Mallotus hokkaidoensis Tanai Hokkaido, Japan Eocene Reference to former two articles Correct?
Tanai, T. 1990 Leaves Mallotus orbiculatus Tanai Hokkaido, Japan Eocene Description, leaf, venation, detailed description margin, no
anatomical details
UIPDD
Tanai, T. 1990 Leaves Mallotus yubariensis Tanai Hokkaido, Japan Eocene Description, leaf, venation, detail margin, no anatomical details Correct?
Vincens, A. 1989 Pollen Macaranga type Burundi, Zaire, Africa Miocene Name mentioned UIPDD
Vincens, A. et al. 2006 Pollen Macaranga type N. Kenya, Africa Oligocene-Miocene Name mentioned UIPDD
Wilf, P. et al. 2007 Leaves Macaranga/Mallotus Laguna del Hunco,
Argentina
Eocene Symposium abstract. Separate photos of leaves, venation only,
no details
UOPDD
Wolfe, J.A 1968 Leaves Macaranga pugetensis Wolfe Washington, U.S.A Eocene Description, leaf fragment, venation only, no details UOPDD
Wolfe, J.A. 1972 Leaves Macaranga type Alaska, U.S.A Palaeocene Leaf fragment, bad condition, venation, no details UOPDD
Yulianto, E. et al. 2004 Pollen Macaranga/Mallotus Batulicin Kalimantan, Indonesia Holocene Name mentioned UIPDD
Yulianto, E. et al. 2004 Pollen Macaranga/Mallotus Pare-Pare Sulawesi, Indonesia Holocene Name mentioned UIPDD
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nectaries), but also because of the state in which the fossils were
preserved. Many fossils were fragmented; missing the apex or the
base of the leaf where the attachment of the petiole is no longer visible;
and even the ones that were found within the present day distribution
cannot be identified reliably (Fig. 1). Therefore, most leaf fossils were
marked as UIPDD or UOPDD in Table 1 and their localities are shown
in yellow and red in Fig. 1.
We cannot provide an extensive analysis of each of the samples in
Table 1. Only the few fossils that showed enough micro-anatomical
details to be reliably identified will be discussed:
−García Massini et al. (2010) described a possible Macaranga
species from the Late Oligocene (27.36±0.11 Ma) of the Plateau
of Ethiopia. The range of characters such as extrafloral nectaries,
disc-shaped glandular hairs and the venation pattern associate
this fossil to both Macaranga and Mallotus; however, due to the
absence of stellate or stellately grouped hairs, the shape of the
leaf and the long hollow petiole the fossil probably has a closer re-
lationship to Macaranga than to Mallotus (Kulju et al., 2007b).
When compared with present day African species of Macaranga
and Mallotus the fossil showed an expected strong resemblance
with Macaranga kilimandscharica Pax (ovate leaf shape with long
acuminate apex, leaf size, position of the extrafloral nectaries,
regularity of the disc-shaped glandular hairs). This is coincidently
a species of Macaranga also present in Ethiopia (Fig. 2).
−Well-preserved Euphorbiaceae leaf macrofossils from the Miocene
(ca. 23 Ma) of Southern New Zealand were reported by Lee et al.
(2010) to be Mallotus-Macaranga. Modern species of Macaranga
and Mallotus, found in Australia and the Pacific, were used to classify
Malloranga fouldenensis D.E.Lee, Bannister, Raine & Coran leaf fossil
(Lee et al., 2010). It is a newly described, monotypic genus, combin-
ing the names of Macaranga and Mallotus, which is indicative of the
problem to assign the fossil to either of them. In the same article (Lee
et al., 2010) the fruits were newly described as Euphorbiotheca
mallotoides D.E.Lee, Bannister, Raine & Coran (genus already
existent, species epithet indicates resemblance with Mallotus). The
pollen (from the fossil inflorescence) was assigned to Nyssapollenites
endobalteus (McIntyre) Kemp & Harris. The Malloranga leaf fossil in
combination with the fruits has the closest resemblance with
Mallotus nesophilus Müll.Arg. (Fig. 3). The similarities are in the
ovate leaf shape and size, venation basally trinerved, position of
the adaxial extrafloral nectaries, random pattern of the abaxial
glandular scale hairs (often in rows in Macaranga) and the type of
gland (compares to Fig. 9 of Fišer Pečnikar et al., 2012, of the re-
lated species M. repandus (Rottler) Müll.Arg.), type and length of the
raceme-like inflorescences (more paniculate in Macaranga), hairy
fruits without spines. The photos (also extra sent on CD-Rom by
Jennifer M.Bannister, co-author of Lee et al., 2010) of the staminate
inflorescences hint at 2-thecate stamens, which confirms the
Mallotus identity. The fruits without spines are typical for section
Philippinenses within Mallotus (Sierra et al., 2005), which includes
M. nesophilus. This section does not contain many species and
M. nesophilus compares best with the fossils availabe. Unfortunately,
it was not possible to use the fossil pollen in order to confirm the
species, since Macaranga and Mallotus pollen are indistinguishable
(see also chapter 3.2).
−Another Euphorbiaceae leaf fossil from the Miocene (ca. 20 Ma) of
New Zealand was described by Mike Pole (Pole, 1996) as being
Macaranga or Mallotus on the basis of its shape, presence of
domatia, a possible pulvinus petiole attachment and venation pat-
tern. This fossil was not as well preserved as the fossils from Lee
et al. (2010) and no digital pictures are available for comparison.
Therefore, further evaluation was not possible and the fossil will
not be used to date the phylogeny.
−Tanai (1989, 1990) described a new species, Mallotus hokkaidoensis
Tanai, from the Middle Eocene (37.2–48.6Ma)ofJapan.Thisfossil
was reported as Mallotus on the basis of lamina shape, petiole, vena-
tion pattern, glandular hairs scattered on the under surface of the
blade and glandular protuberances on the margin formed by the
ends of the tertiary venation. It must be mentioned, that in earlier
literature (Tanai, 1968–70), the glands on the underside of the
blade of Mallotus hokkaidoensis were seemingly not observed so
that identification was still a problem, and Tanai (1968–1970)
even compared it with Ficus columboides Endo and F. ezoensis
Endo (Table 2 shows that leaf venation patterns of some Ficus
species (Moraceae) look similar to those of Macaranga and
Mallotus; see Discussion). Mallotus hokkaidoensis shows close
similarities to the modern species M. philippensis (Lam.) Müll.Arg.,
M. japonicus Müll.Arg. and M. paniculatus (Lam.) Müll.Arg., but
mostly to M. japonicus (Tanai, 1990) because of the thin quaternary
veins and the shape of the leaf lamina (ovate to trilobed). It was also
stated that this new species closely resembles the fossil species
Mallotus oregonensis Sanborn (Sanborn, 1937)andM. riparius
MacGinitie (MacGinitie, 1941), although the secondary venation is
somewhat different.
−Mallotus yabariensis Tanai (Tanai, 1990) from the Eocene (37.2–
48.6 Ma) of Japan, owes its classification to its lamina shape, size,
margin, venation pattern (secondary venation ending in obtuse
teeth from a large marginal loop) and punctate glands scattered
on the under surface of the blade as dark spots. Mallotus yabariensis
closely resembles the modern species M. tenuifolius Pax (Tanai,
1990), because of the protuberances along the margin, trilobed
lamina and well-developed tertiary and slender transversed quater-
nary veins (wording after Tanai, 1990). This fossil and the former
one discussed date from the same period and can be used as calibra-
tion points for dating a phylogeny.
The geographic distinction between the classes UOPDD and UIPDD
is not always sharp. Some of the fossils classified as UOPDD could al-
most be categorized as UIPDD or possibly being Macaranga or Mallotus
Fig. 2. Upper (left) and lower (right) surface of leaves of Macaranga kilimandscharica
Pax. Leaves of this species resemble the fossil leaves pictured by García Massini et al.
(2010) closest in anatomical details, compare with their Fig. 13.3-6. [left: Brass
17294, right: Maas Geesteranus 5641, both L].
108 M. Nucete et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 353–355 (2012) 104–115

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species: Fossils of Macaranga peltata (Roxb.) Müll.Arg. (Akhmetiev and
Vikulin, 1995)andMallotus rjufflei Ablaev et Ig. Vassiljev (Ablaev and
Vassiljev, 1994) found along the eastern coast of Russia are said to be
from the Oligocene (23.3–33.9 Ma). During that time, Japan, due to
the positioning of the tectonic plates, was very close to mainland
Russia, making migration possible (Fig. 8). Present day Japan has
species of Mallotus as part of its flora, and fossils found and identified
as Mallotus marked as UIPDD may well be correctly identified (Fig. 1).
3.2. Pollen
The best fossil pollen available was the Nyssapollenites endobalteus
pollen described by Lee et al. (2010). Raymond van der Ham indicated
that the fossil pollen found in New Zealand (Lee et al., 2010) could not
be reliably identified as Macaranga or Mallotus. The pollen grains of
both genera look similar, and similar pollen occurs throughout the
acalyphoid euphorb clade Hancea (formerly Cordemoya s.l.), Macaranga
and Mallotus (Takahashi et al., 2000; Nowicke and Takahashi, 2002).
The fossil pollen clearly belongs in this clade, but pollen itself is not
suitable to narrow the identification down to a genus, let alone a
species (Jennifer Bannister and Raymond van der Ham, pers. comm.).
All other fossil pollen (Table 1) was of less quality than the pollen
shown by Lee et al. (2010) or pollen was only mentioned, therefore
these samples are not evaluated here, because identification is impos-
sible due to the generality of the pollen characters.
3.3. Seeds
The pictures of the only seed fossils (Mai, 2001) were analyzed by
Wolfgang Stuppy. Three seeds were discussed and identified as
Mallotus tuberculatus Mai (Mai, 2001). One of the fossil seeds described
is campylotropous (Mai, 2001,histable12,figure 11). Euphorbiaceae
(almost) invariably have anatropous seeds (Stuppy, 1995). It is there-
fore highly unlikely that this illustrated seed belongs to either Mallotus
or Macaranga. The other seeds discussed in that paper (Mai, 2001,his
table 12, figures 8–10) show typical Euphorbiaceae characters such as
an ovoid shape, smooth surface, and, most distinctly, a sclerenchyma-
tous layer in the seed coat that appears to consist of a single layer of
long, diagonal palissadal cells. The latter is the hallmark character for
Euphorbiaceae (s.s.) seeds, but only at family level (Stuppy, 1995).
Unfortunately, most Euphorbiaceae seeds display too little variation
in characters to allow even identification to genus level (few excep-
tions permitted, e.g., Ricinus,Hevea). Two of the fossil seeds described
Table 2
List of genera from various families resembling the venation patterns in Macaranga and
Mallotus. Between brackets former family names often still present in literature.
Family Genus
Cornaceae (Alangiaceae) Alangium (some)
Bignoniaceae Catalpa
Datiscaceae Octomeles
Elaeocarpaceae Sloanea (some)
Euphorbeaceae Endospermum
Melanolepsis
Achariaceae (Flacourtiaceae) Trichadenia
Hernandiaceae Hernandia
Malvaceae Hibiscus (some)
Thespesia
Malvaceae (Sterculiaceae) Byttneria (some)
Firmiana
Heritiera (some)
Kleinhovia
Pterospermum
Sterculia
Malvaceae (Tiliaceae) Brownlowia
Trichospermum
Menispermaceae Coscinium
Moraceae Ficus (some)
Paulowniaceae (Scrophulariaceae) Paulownia
Verbenaceae Clerodendrum (some)
Fig. 3. Upper (left) and lower (right) surface of leaves of Mallotus nesophilus Müll.Arg., fruit (center) c. 5 mm. This species most closely resembles the fossil material Malloranga
fouldenensis D.E Lee, Bannister, I.J. Raine & Coran (Lee et al., 2010)[Gray 6646, L].
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by Mai (2001) are clearly Euphorbiaceae seeds, but Macaranga and
Mallotus belong to the group of genera with the very general type of
seeds, thus no identification up to genus level is possible.
3.4. Wood
Elisabeth Wheeler (co-author of Gregory et al., 2009) evaluated
the fossilized wood (Lakhanpal and Dayal, 1963; Kramer, 1974;
Prakash and Tripathi, 1975; Roy and Ghosh, 1982; Guleria and
Srivastava, 2001) and according to her the wood characteristics
found are too general to identify reliably even to family level. In
fact, identification is also impossible because the wood of extant
Macaranga and Mallotus is also not characteristic.
4. Discussion
We are confident that we collected most literature describing
Macaranga and Mallotus fossils. Especially the modern articles are im-
portant, because the authors have applied more modern research
methods now available, which revealed the micro-anatomical charac-
ters we needed to be certain abouta possible identification. Most fossils
referred to Macaranga or Mallotus were imprints of leaf fragments.
These were usually described as having an ovate or elliptic shape with
a round base, entire (or serrate) margin and an acute to apiculate
apex. The venation, generally deemed most important for identification,
is usually described as either pinnate or basally palmate or 3-nerved
(and higher up pinnate) with camptodromous or acrodromous secondary
veins (nerves curving upwards towards the leaf margin and usually
joining in a series of conspicuous marginal loops), scalariform (ladder-
like) tertiaries and reticulate quaternaries. Macaranga and Mallotus
species are very variable in their venation pattern and certainly cover
all venation patterns found in the fossils, especially when fossil frag-
ments are considered. However, the leaf and venation description is
not unique to Macaranga and Mallotus only, other genera easily fitthe
description too. Figs. 4 and 5 show some examples and Table 2 contains
a non-exhaustive list of possibilities, compiled by Max van Balgooy, of
other Malesian contemporary genera (Euphorbiaceae and other
families) with similar venation patterns. It is likely that the African
and South American tropical floras also contain numerous taxa from
different families with the same general type of venation and leaf
shape (especially when only leaf fragments are considered). Another
confirmation that leaf shape and venation does not show sufficient de-
tail for a correct identification is that a fossil, previously described as
Mallotus comstockii Sanborn (Sanborn, 1937), was renamed Platanus
comstockii (Sanborn) J.A.Wolfe (Wolfe, 1977) and was thus referred to
the completely different, non-related family Platanaceae (order Proteales;
Euphorbiaceae order Malphigiales). It appears that leaves with a
camptodromous or acrodromous venation have been ascribed to
Macaranga and Mallotus so many times that new fossils are automatically
referredtothesetwogenera.Herewehavedemonstratedthatthevena-
tion pattern by itself is not sufficient to identify fossils as Macaranga or
Mallotus.
Fig. 1 is a present day map (Blakey, 2011) showing the localities of
the fossils. The different shapes indicate the different genera; the
Fig. 5. Similarities in venation patterns between taxa from various families. Secondary veins looping and connecting near the margin, tertiary veins scalariform: A. Microcos hirsuta
(Korth.) Burret (Tiliaceae); B. Mallotus philippensis (Lam.) Müll.Arg. (Euphorbiaceae) [A: SAN (Amin et al.) 107147,B:Waas 758, both L].
Fig. 4. Similarity in leaf blade shape and venation between taxa in various families: A. Alangium rotundifolium (Hassk.) Bloemb. (Alangiaceae/Cornaceae); B. Mallotus tiliifolius
(Blume) Müll.Arg. (Euphorbiaceae) [Korthals s.n., L 901.169-341; KL (Teo & Pachiappan) 3104; both L].
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colours indicate the degrees of identification reliability (green is reli-
able) and presence within or outside the present day distributions of
Macaranga (black line) and Mallotus (white line). As you can see from
Fig. 1, most of the fossils considered in this paper have been found in
India. Modern species of Macaranga and Mallotus are native to India.
This makes it well possible that these fossils were correctly identified.
However, the fossils found in India are badly preserved and no
micro-anatomical details were described, nor was it not possible to
Fig. 6. Locality sites of the fossils in their geological time (Blakey, 2011). Different shapes indicate the genera and the different colours represent the degrees of reliability: Δ=
Macaranga fossils; ○=Mallotus fossils; □=Macaranga or Mallotus fossil; Green=reliable fossil; Yellow = Unidentifiable withIn the Present Day Distribution (UIPDD); Red =
Unidentifiable Outside the Present Day Distribution (UOPDD). Maastrichtian and Palaeocene (55.8–70.6 Ma). Eocene (33.9–55.8 Ma). Oligocene (23.03–33.09 Ma). Miocene
(5.332–23.03 Ma). Holocene, Pleistocene, Pliocene (Present −5.332 Ma). Maastrichtian and Palaeocene (55.8–70.6 Ma).
Fig. 7. Locality sites of the fossils intheir geological time (Blakey, 2011). Different shapes indicate the genera and thedifferent colours represent the degreesof reliability: Δ=Macaranga
fossils;○=Mallotus fossil s; □=Macaranga or Mallotus fossil; Green= reliable fossil; Yellow=UnidentifiablewithIn the Present Day Distribution (UIPDD); Red= Unidentifiable Outside
the Present Day Distribution (UOPDD). Maastrichtian and Palaeocene (55.8–70.6 Ma). Eocene (33.9–55.8 Ma). Oligocene (23.03–33.09 Ma). Miocene (5.332–23.03 Ma). Holocene,
Pleistocene, Pliocene (Present−5.332 Ma). Eocene (33.9–55.8 Ma).
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acquire any additional pictures or information. Therefore, based on
the publications these fossils were identified as UIPDD.
Historical biogeographic methods use the distributions of contem-
porary species to induce the possible distributions of ancestral species.
The major disadvantage is that it is impossible to obtain ancestral dis-
tributions outside the present day distributions (white and black
lines in Fig. 1). Fossils might give an indication about different distribu-
tions in former geological times. In the remaining part of the discussion
Fig. 9. Locality sites of the fossils in their geological time (Blakey, 2011). Different shapes indicate the genera and the different colours represent the degrees of reliability: Δ=
Macaranga fossils; ○=Mallotus fossils; □=Macaranga or Mallotus fossil; Green=reliable fossil; Yellow = Unidentifiable withIn the Present Day Distribution (UIPDD); Red =
Unidentifiable Outside the Present Day Distribution (UOPDD). Maastrichtian and Palaeocene (55.8–70.6 Ma). Eocene (33.9–55.8 Ma). Oligocene (23.03–33.09 Ma). Miocene
(5.332–23.03 Ma). Holocene, Pleistocene, Pliocene (Present −5.332 Ma). Miocene (5.332–23.03 Ma).
Fig. 8. Locality sites of the fossils intheir geological time (Blakey, 2011). Different shapes indicate the genera and thedifferent colours represent the degreesof reliability: Δ=Macaranga
fossils;○=Mallotus fossil s; □=Macaranga or Mallotus fossil; Green= reliable fossil; Yellow=UnidentifiablewithIn the Present Day Distribution (UIPDD); Red= Unidentifiable Outside
the Present Day Distribution (UOPDD). Maastrichtian and Palaeocene (55.8–70.6 Ma). Eocene (33.9–55.8 Ma). Oligocene (23.03–33.09 Ma). Miocene (5.332–23.03 Ma). Holocene,
Pleistocene, Pliocene (Present−5.332 Ma). Oligocene (23.03–33.09 Ma).
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all fossils will be considered as possible Macaranga and/or Mallotus fos-
sils and their geographical position (relative to the positions of the tec-
tonic plates) will be evaluated in a possible historical biogeographical
scenario. This is another test of the identification status of the fossils.
When comparing Figs. 6–10 (Palaeontological maps of the fossils
found from the Maastrichtian to the Holocene) it is clear that most
possible Macaranga and Mallotus fossils are from the Holocene, Pleisto-
cene, Pliocene (Fig. 10)andMiocene(Fig. 9). This supports the idea of
recent speciation (Kulju et al., 2007b; Sierra et al., 2010). Also, it can be
seen from these maps that most of the fossils reported to be much
older, were found far outside the present day distribution in North
and South America and Europe. It is tempting to think that both sister
genera originated in the western hemisphere, but unfortunately there
are no micro-anatomical details preserved in these fossils to support
this idea. The basal species in the phylogeny of Macaranga (Kulju et
al., 2007b)andMallotus (Kulju et al., 2007b; Sierra et al., 2010)are
from Borneo, which presents a completely different view (but not
necessarily correct as migration may have occurred). Based on the
chronogram in Davis et al. (2005) an age of c. 90 Ma is expected for
the ancestral species of Macaranga and Mallotus.
The palaeontological map of the Maastrichtian (Late Cretaceous;
Fig. 6) shows the possible localities of the two oldest fossils. These
are far apart and one is even far outside the present day distribution
of Macaranga and Mallotus. The Indian Mallotus fossil is thought to
be from the Maastrichtian and the Alaskan Macaranga fossil from
the Palaeocene. Thus, the Indian fossil is older, which may imply a
possible center of origin for perhaps both sister taxa in India. But as
no typical characters were visible on the Indian (nor the Alaskan) fos-
sil, it is impossible to draw firm conclusions about an area of origin,
also because in the next period, the Eocene (Fig. 7), no fossils from
India are known. Finding an area of origin based on fossils is difficult,
because Macaranga and Mallotus mainly occur in the everwet tropical
regions, areas in which fossilization is poor. Most fossils are found at
the margins of the present day distributions or outside those distribu-
tions, all areas with better possibilities for fossilization.
Most fossils from the Eocene (Fig. 7) were found far outside the
present day distribution of Macaranga and Mallotus. The Alaskan
Mallotus comstockii fossil, that was later re-identified as Platanus
comstockii, is one of them. There are two fossils on this map that
were found within the present day distribution, Australia and Japan.
The Japanese fossil, Mallotus hokkaidoensis, is the oldest fossil from
our reliable list. At this time India was already attached to the mainland
making it possible for the genera to have spread. Unfortunately,
Australia was still far away and, therefore, it is illogical to assume
that the genera could have originated in India and spread to Australia.
The fossils in this map are so spread out that it is impossible to specu-
late on the area of origin of the genera. Moreover, it is even unlikely
that any of the fossils is really Macaranga or Mallotus except the
Japanese ones, especially if the genera originated in S.E. Asia.
In the Oligocene (Fig. 8) we can see the second oldest of the reliable
fossils. It was found in Africa. In this map there are no fossils present in
India. The fossils are spread through Africa, Europe, Asia and Australia.
Unfortunately, the European fossil could not clearly be identified as
Macaranga or Mallotus. Dispersal between Africa, India and Asia was
probably relatively common at different geological times and in both
directions (Li et al., 2009). Most of the mentioned genera, like
Macaranga and Mallotus, are not known for long-range dispersal, thus
the climate must have been warmer and wetter with more forest pres-
ent, allowing dispersal either along the southern coast of Asia or along
island chains between India and Africa/Madagascar (e.g., Kulju et al.,
2007a).
In Figs. 9 and 10 we can see that most fossils from the Miocene and
younger were found in India and inside the present day distribution.
Probably, a number of these fossils may have been correctly identified,
but that can not be confirmed. The reliable Mallotus fossil from New
Zealand is present in the Miocene map (Fig. 9); as at that time New
Zealand and Australia had moved closer to the mainland. It seems plau-
sible that the spread could have occurred from India eastwards to Asia
and then to Australasia. Probably, climatic conditions were warmer,
allowing Mallotus to reach New Zealand. Mallotus probably disperses
Fig. 10. Locality sites of the fossils in their geological time (Blakey, 2011). Different shapes indicate the genera and the different colours represent the degrees of reliability: Δ=
Macaranga fossils; ○=Mallotus fossils; □=Macaranga or Mallotus fossil; Green=reliable fossil; Yellow = Unidentifiable withIn the Present Day Distribution (UIPDD); Red =
Unidentifiable Outside the Present Day Distribution (UOPDD). Maastrichtian and Palaeocene (55.8–70.6 Ma). Eocene (33.9–55.8 Ma). Oligocene (23.03–33.09 Ma). Miocene
(5.332–23.03 Ma). Holocene, Pleistocene, Pliocene (Present −5.332 Ma). Holocene, Pleistocene, Pliocene (Present –5.332 Ma).
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better than Macaranga or can adapt better to more temperate climates,
because nowadays, Mallotus is present in Japan and northern India,
Macaranga is not. This is another confirmation of our idea that
Malloranga fouldenensis might be a Mallotus species.
The fossils do not facilitate a choice of area of origin for both gen-
era. Even the oldest fossil in India is not found in an everwet tropical
climate. It is very likely that the origin must have been in a wet
tropical climate in Asia, India, and/or Africa as by far the majority of
species occurs in these areas. Extensions of the distributions to
Europe may have happened, just as it happened to New Zealand,
but a palaeontological presence in the Americas seems highly
unlikely.
When looking for reliable fossils to date a phylogeny then the
following procedure seems best practice:
−Cooperate with a palaeontologist, in our case the second author
and her network of experts, to get an overview of taxa described
and literature available. Also use the Internet and references in
literature about fossils.
−Collect the literature and obtain a first impression of the data
available. If the data seem insufficient ask more information
(e.g., digital photos) from the authors, many are willing to comply.
−Establish which characters are important for a reliable identifica-
tion and if you feel inadequate to evaluate a certain fossil contact
experts (as we did for wood, pollen and seeds).
−Sort the fossils mentioned in literature as adequately or inade-
quately identified. When deemed adequate then try to link them
via morphology to contemporary species.
−See if a historical biogeographic scenario is possible.
Acknowledgments
Dr. Raymond W.J.M. van der Ham, Dr. Max M.J. van Balgooy, Dr.
Elisabeth A. Wheeler and Dr. Wolfgang Stuppy are thanked for sharing
their knowledge and expertise. Dr. Ronald Blakey for providing the
palaeontological maps and Jennifer M. Bannister from the University
of Otago, Diane M. Erwin from the University of California Museum of
Paleontology, Dr. Bonnie F. Jacobs Chair of the Environmental Science
Program at SMU, Dr. Barbara Mohr at the Museum für Naturkunde;
Leibniz-Institut für Evolutions- und Biodiversitätsforschung and the
Humboldt-Universität zu Berlin, Dr. Kazuhiko Uemura from the
National Science Museum, Tokyo and Dr. Peter Wilf at the University
of Pennsylvania for sending high quality pictures for further analysis
of the fossils mentioned in this paper and for being readily available.
Dr. Niels Raes for knowledge on editing maps and Mr. Ben Kieft for
making photos. The two reviewers are thanked for their improvements
to the manuscript.
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