# Carya nux-taurinensis comb. nov., a fossil-species incorporating Carya globosa (Juglandaceae)

Edoardo Martinetto, Dario De Franceschi, Cedric del Rio, Steven R Manchester

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Edoardo Martinetto, Dario De Franceschi, Cedric del Rio, Steven R Manchester. Carya nux-taurinensis comb. nov., a fossil-species incorporating Carya globosa (Juglandaceae). Taxon, the journal of the International Association for Plant Taxonomy, In press, 10.1002/tax.70065. hal-05339092

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RESEARCH ARTICLE

*Carya nux-taurinensis* comb. nov., a fossil-species incorporating *Carya globosa* (Juglandaceae)

Edoardo Martinetto,¹ Dario De Franceschi,² Cedric Del Rio² & Steven R. Manchester³

1 Dipartimento di Scienze della Terra, Università di Torino, Via Valperga Caluso 35, 10125 Torino, Italy

2 CR2P – Centre de Recherche en Paléontologie – Paris, MNHN – Sorbonne Université – CNRS, 43 Rue Buffon, 75005 Paris, France

3 Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, U.S.A.

Address for correspondence: Edoardo Martinetto, edoardo.martinetto@unito.it

DOI: https://doi.org/10.1002/tax.70065

**Abstract** The type material of the almost forgotten fossil-species *Juglans nux-taurinensis* is newly analysed, a lectotype is designated, and new conspecific material detected at the Italian type locality La Morra is described. We show that *Juglans nux-taurinensis* must be reassigned to *Carya* because the locule cast of the nutshell shows two inner symmetrical longitudinal ribs protruding into the locule, which are characteristic of *Carya* and missing in *Juglans*. The new materials from La Morra clearly demonstrate that the nutshell morphology matches that of *Carya globosa*, a name that we consider to be a younger heterotypic synonym of *Carya nux-taurinensis* comb. nov.

**Keywords** Europe; fossil-species; Neogene; nomenclature; nuts; palaeobotany; synonym

**Supporting Information** may be found online in the Supporting Information section at the end of the article.

■ INTRODUCTION

*Juglans nux-taurinensis* Brongn. is the name of a fossil-
species (see Art. 1.2 of the ICN, Turland & al., 2018) of Ju-
glandaceae based on a locule cast and an external mold of nut-
shells, whose origin was indicated as the “hills around Turin”
(Brongniart, 1822). Fossil fruits of Juglandaceae are rather
abundant in the Cenozoic of Europe and have been intensively
studied (e.g., Kirchheimer, 1957; Manchester, 1987;
Mai, 1981; Deng & al., 2024), but this species of Brongniart
(Fig. 1B,D) has been almost forgotten, possibly due to lack
of detailed information and adequate photographic
documentation.

Brongniart's collection is housed at Muséum national d'Histoire naturelle (MNHN) of Paris. A search carried out by one of us (D.D.F.) allowed us to locate the original specimens classified as *Juglans nux-taurinensis*, bearing the locality indication “La Morra” (Fig. 1A,C). This site is in the region of Turin (Piemonte, NW Italy), but not in the “hills around Turin” (50 km apart), which suggests that Brongniart (1822) had only a vague idea about the origin of the fossils. Nuts originating from La Morra were known since the eighteenth century and the first citation of a fossil nut from the site is probably that of Allioni (1757: “*Carpolithus juglandis*”, still available in the Museo Geologico Sperimentale of Giaveno, Italy, MGSG: Martinetto & al., 2023). This proves that

fossil nuts from La Morra were known well before the starting
point for palaeobotanical nomenclature (31 December 1820:
ICN, Art. 13.1, thus invalidating Allioni's name).

In this paper, our aim is to review and revise the fossil-
species *Juglans nux-taurinensis* Brongn. based on new fossil
discoveries and updates in the knowledge of fruit morphology
of extant Juglandaceae. We conclude that *J. nux-taurinensis*
must be recombined to *Carya*. Additionally, we consider
*C. globosa* (R.Ludw.) Mädler (Fig. 1F–H) to be a synonym
of *Carya nux-taurinensis* comb. nov., indicating that this spe-
cies was widely distributed in the Neogene of Europe.

■ MATERIALS AND METHODS

We have examined 10 specimens formerly attributed to *Juglans nux-taurinensis*. The five specimens from La Morra at the MNHN probably correspond to two sets: One from an unknown collector, which is represented by two specimens (MNHN.F.3010.1, MNHN.F.3010.3), and another set of three specimens (MNHN.F.3010.2, MNHN.F.3010.4, MNHN.F.3010.5: Figs. 1E, 2A) sent by Eugène (Eugenio) Sismonda in 1834 (according to the labels). The second set of specimens does not correspond to the specimens figured by Sismonda (1865: pl. 29, figs. 11–13), which were housed in Turin, at the “Regio Museo Geologico” (Peola, 1895) and are most
<figure><img src="image_3.png" /><figcaption>**Fig. 1.** A–E, *Carya nux-taurinensis* comb. nov. from the Late Miocene of La Morra, Piemonte region, NW Italy. A, Lectotype (designated in this paper), locule cast (MNHN.F.3010.1, view parallel to the primary septum) corresponding to the morphology of the inner surface of a nutshell; B, Illustration by Brongniart (1822: pl. VI, fig. 6b) associated to the valid publication of the name *Juglans nux-taurinensis*, considered by us to represent the specimen of Fig. 1A; C, Syntype MNHN.F.3010.3, mold recording the external morphology of a nutshell different from the previous one, but assigned both by Brongniart (1822) and by us to the same species; D, Illustration by Brongniart (1822: pl. VI, fig. 6a) considered by us to represent the specimen of Fig. 1C; E, MNHN.F.3010.4 (“Sismonda collection”), external mold of a nutshell with prominent grooves. F–H, Drawings of the holotype of *Carya globosa* (here considered a synonym of *C. nux-taurinensis*) from the Pliocene of Dorheim, Germany (Ludwig, 1857) showing the external surface of the organically preserved nutshell (F) and the internal morphology of the two halves (G & H) resulting from nutshell’s splitting along the plane of the secondary septum; primary septum visible in the centre of the nutshell (H). — Scale bars: 5 mm.</figcaption></figure>

<figure><img src="image_4.png" /><figcaption>**Fig. 2.** *Carya nux-taurinensis* comb. nov. Locule casts of complete, closed nutshells from La Morra, reproducing the morphology of the inner surface; A, Specimen MNHN.F.3010.5, "Sismonda collection"; B, Specimen stored at the Museo Civico Craveri of Bra (MCCB-04208), representative fossil for the Piemonte region selected by the Italian Palaeontological Society. — Scale bars: 5 mm.</figcaption></figure>
probably lost (Martinetto & al., 2023). Both sets of MNHN materials have been merged into the “Sismonda collection”, probably because of the same locality mentioned in the labels, and same plant fossil taxon. This must have been done in 1835, when the inventory catalogue of MNHN started (nr. MNHN.F.3010 is from that period). Additionally, we examined five other specimens from the same locality and species: Four locule casts deposited in the Craveri Museum of Bra (in particular one: MCCB-04208, Fig. 2B) and a locule

cast at Museum für Naturkunde of Berlin (MB: Fig. 3J). We also studied a nutshell mold at MGSG (Fig. 3I) and, at the Civic Museum Eusebio of Alba (MCEA), an external mold of a nutshell (MCEA-P05043: Fig. 3K) from Case Missione (close to the town of Alba, Late Miocene: Martinetto & al., 2023: 28, fig. 8D), which appeared similar to some of the above-mentioned specimens from La Morra.

In December 2024 we detected the most probable site of origin of Brongniart's fossils, close to the hamlet Croera

<figure><img src="image_5.png" /><figcaption>**Fig. 3.** *Carya nux-taurinensis* comb. nov. from the Late Miocene of the Piemonte region, NW Italy; all from La Morra, except K. A, Aspect of the Late Miocene fossil-bearing sandstones of La Morra with the external mold of a complete nutshell showing striking grooves (indicated by the finger), MRSN-P/345-CCN8670; B, Apical part of the external mold of a complete nutshell with striking grooves, MRSN-P/345-CCN8671a; C, Remaining part (basal) of the external mold shown in Fig. 2B, MRSN-P/345-CCN8671b; D, Locule cast (view parallel to the primary septum) corresponding to the morphology of the inner surface of a halved nutshell, MRSN-P/345-CCN8672; E, Partially exposed locule cast of a nutshell (basal part), MRSN-P/345-CCN8673. F–H, MRSN-P/345-CCN8674, specimen combining the external mold of a half nutshell and the associated, partially exposed locule cast in basal (F) and oblique lateral view (G & H); notice the undulated outline recording the external morphology of the nutshell with striking grooves (arrow in F) and ridges (arrow in H); the two basal lobes of the locule cast, separated by the secondary septum, are indicated by the arrows in G. I, External mold of the basal part of a nutshell with striking grooves and three ridges deriving from post-burial mineral infilling, that could correspond to the three fissures at right angles that may occur in fossil nutshells, as it is visible in Fig. 4C; from the Allioni collection (A.C. 1370) at the MGSG, collected at La Morra in the 18th century (Allioni, 1757); J, Partially exposed locule cast of a nutshell; the arrow points to one of the two basal lobes separated by the secondary septum, MB.Pb.2014/0643; K, External mold of a nutshell with striking grooves; from Case Missione, close to the town of Alba, Late Miocene, Civic Museum Eusebio of Alba (MCEA-P05043). — Scale bars: 5 mm.</figcaption></figure>
of La Morra, where a marine geological formation named “La Morra Unit” by Ghielmi & al. (2019) crops out on a steep slope. The age of these sediments is indicated as early Messinian (Late Miocene, ca. 7.0–5.9 Ma) by Ghielmi & al. (2019), a chronostratigraphic attribution also supported by other geological data from the La Morra area (Dela Pierre & al., 2011). A small boulder, mainly made up of conglomerates, contained a lens of sandstone that yielded a locule cast of a halved nut (Fig. 3D) and three complete (still empty) external molds of nutshells (Fig. 3A–C), all of which corresponded to the morphologies described by Brongniart (1822) for *Juglans nux-taurinensis* (*C. nux-taurinensis* comb. nov.). The sandstone in which the new fossil nuts are preserved has the same appearance as the matrix of the fossils conserved at MNHN, which is characterised by early diagenetic cementation (cite?) and decay of the organic matter, whose original presence is rarely indicated by dusty black remains (Fig. 3H) and, nearly always, by reddish patinas that coat the external molds of abundant plant remains, which left empty cavities (Fig. 3A). The nutshells are often preserved as external molds (Figs. 1C,E, 3A–C,I) of halved or complete specimens. The

internal surfaces of tightly closed nutshells were not preserved and the successive decay of the organic matter left empty globose cavities (Fig. 3A–C), whereas in halved nutshells the internal morphology was reproduced on the sediment, thanks to the cementation of the sandstone before the decay of the nutshell's organic matter (Fig. 3D–H,J). The new materials from the type locality La Morra (MRSN-P/345-CCN8670–8674: Fig. 3A–H) were deposited at the CENOFITA collection (acronym MRSN-P/345-CCN), managed in synergy between the Turin University and the Regional Museum of Natural Sciences of Turin (MRSN).

Since we noticed a similarity of the fossil nuts from La Morra to *Carya globosa* (Mädler, 1939; Mai, 1981; Manchester, 1987), we checked at MB the holotype (Fig. 1F–H) figured by Ludwig (1857) and the Italian material assigned to this last fossil-species, stored in the CENOFITA collection (Martinetto, 2015). In particular, a well-preserved nut from Canton Iuli, very similar to the holotype of *C. globosa*, was analysed by micro-CT scanning (Fig. 4A–F) in order to obtain data useful to compare the internal morphology to that of the nuts attributed to *Juglans nux-taurinensis*.

<figure><img src="image_6.png" /><figcaption>**Fig. 4.** Micro-CT scanning of a fossil nut and extant species for comparison. **A–F**, Fossil *Carya nux-taurinensis* comb. nov., MRSN-P/345-CCN8675, virtual reconstruction of different parts of an organically preserved nutshell formerly assigned to *C. globosa* (Martinetto, 2015) from the Pliocene site Canton Iuli, Piemonte region, NW Italy: **A**, Locule cast, view parallel to the primary septum, which corresponds to the morphology of the inner surface of the nutshell; **B**, External nutshell surface, lateral view; **C**, External surface, basal view, notice the fissures in the basal part; **D**, Locule cast, basal view; **E**, Locule cast, view orthogonal to the primary septum; **F**, Cross section of the nutshell with scanty remains of the seed coat inside, notice the contact between seed coat and nutshell's inner surface. The red and green lines indicate the position of two virtual sections at right angles, not shown in this paper. **G**, Extant *C. pallida*, FLAS76003 (J.D. Ray Jr 7292, Mississippi, U.S.A.), virtual cast of the seed locule, which corresponds to the morphology of the inner surface of the nutshell (Deng & al., 2024), because the seed is tightly included in the nutshell; **H**, Extant *C. tomentosa*, FLAS88222 (D. Demaree 43185, Arkansas, U.S.A.), virtual locule cast; **I**, Extant *C. ovata*, UF872 (D. Dilcher s.n., Indiana, U.S.A.), virtual locule cast; **J**, Extant *Juglans regia*, cross section of a nutshell with the seed inside, notice the gap between the seed and nutshell's inner surface. — Scale bars: 5 mm.</figcaption></figure>
For the interpretation of the morphology of the fossils and their taxonomic significance, we studied (suppl. Table S1) the nuts of eight extant species of *Carya* (three shown in Fig. 4G–I) and eight of *Juglans* (only one shown in Fig. 4J), by micro-CT scanning using the same specimens already illustrated by Deng & al. (2024) and Manchester & al. (2024: fig. 9). These originate either from the palaeobotanical collection (UF) or the herbarium (FLAS) of the Florida Museum of Natural History at the University of Florida, Gainesville. Tiff stacks, herbarium voucher data, and scanning parameters for the extant nuts of *Carya* and *Juglans* that were micro-CT scanned are archived at *morphosource.org* (project: “Juglandaceae nuts”).

The new combination and the lectotypification were registered with PFN numbers in the Plant Fossil Names Registry (https://www.plantfossilnames.org/), hosted and operated by the National Museum, Prague for the International Organisation of Palaeobotany (IOP).

## ■ RESULTS AND DISCUSSION

The species *Juglans nux-taurinensis* was described by Brongniart in 1822. The numbers MNHN.F.3010.1 (Fig. 1A) and MNHN.F.3010.3 (Fig. 1C) correspond to Brongniart's (1822: pl. VI) figures 6b (Fig. 1B) and 6a (Fig. 1D), respectively. Brongniart described and figured the locule cast of a halved nutshell and an external nutshell mold, seemingly implying both were parts of the same specimen. Brongniart (1822: 323, pl. VI, figs. 6a, 6b) validly published the name *J. nux-taurinensis* with a very short diagnosis (“nuce globosa, sulcata, rugosa, seminis lobis approximatis simplicibus”). This description and the associated drawings are sufficiently informative to be sure that the locule cast of a halved nutshell MNHN.F.3010.1 and the external mold MNHN.F.3010.3 are the specimens of *J. nux-taurinensis* described in the protologue. Even if their origin from a single nut was not so explicitly indicated, Brongniart (1822) definitely considered the cast MNHN.F.3010.1 to be conspecific to the external mold MNHN.F.3010.3, and in the protologue he described the nut as characterised by a very rough external surface (Brongniart, 1822). However, our observations indicate that the two samples illustrated by Brongniart (1822) on plate VI, figs. 6a and 6b, are from two different specimens. Even if they came from the same locality (same collection, it seems: Same lithology, labels, patina, etc.), they are not complementary samples (not adjustable to be put together).

It seems that Brongniart (1822) in his plate VI, fig. 6b (Fig. 1B) did not depict (as was usual at that time) the sediment around the sample, but the outline of the locule cast presented on the plate coincides with that preserved on the sediment. An artificial cast (wax and plaster) of MNHN.F.3010.1 stored at MNHN shows a good correspondence with Brongniart's fig. 6b. The second set of samples of the “Sismonda collection” (MNHN.F.3010.2, 4, 5) was not cited by Brongniart (1822) and, consequently, only MNHN.F.3010.1 and MNHN.F.3010.3 are the syntypes of *Juglans nux-

*taurinensis*. Since Brongniart (1822) figured two specimens, the holotype cannot be designated with certainty, and we designate as lectotype the specimen MNHN.F.3010.1 (Fig. 1A), housed at Muséum national d'Histoire naturelle of Paris, considered to be the fossil drawn by Brongniart (1822) in his plate VI, fig. 6b.

The availability of the original specimens of Brongniart (1822) allowed us to carry out an updated analysis of the morphology in light of the advances in the knowledge of the Juglandaceae fruit and seed morphology (Manchester, 1987; Mai, 1981; Deng & al., 2024). After careful consideration of the morphological characters, we conclude that *Juglans nux-taurinensis* must be recombined into *Carya* Nutt. because the locule cast of the nutshell shows two inner symmetrical longitudinal ribs protruding into the locule (Fig. 1A), which are characteristic of *Carya* and missing in *Juglans* L. (Manchester & al., 2024). *Juglans* is represented in the late Cenozoic of Italy by a rather widespread and well-known species, *J. bergomensis* (Bals.-Criv.) A.Massal. (Martinetto & al., 2015), with an internal nutshell (and seed) morphology particularly similar to that of the extant *J. cinerea* L. (Deng & al., 2024: fig. 1S–U), and is quite distinct from the morphology of *Juglans nux-taurinensis*.

Therefore, we formally propose the new combination *Carya nux-taurinensis* comb. nov. Our recent field work at La Morra yielded a halved specimen (MRSN-P/345-CCN8674) with combined cast and mold preservation of the nutshell, respectively similar to MNHN.F.3010.1 (lectotype) and MNHN.F.3010.3 (remaining syntype), proving that the two morphologies are associated (Fig. 3F–H). Three additional specimens from the locality La Morra completed our understanding of the nutshell morphology of *C. nux-taurinensis*: Two locule casts of whole nutshells with scanty petrified parts, MNHN.F.3010.5 and MCCB-04208 (Fig. 2A,B), and the external mold MNHN.F.3010.4 (Fig. 1E). When we compared the internal nutshell morphology of the six above-mentioned fossils to analogous parts of the extant relatives (*Carya* and *Juglans* species; Fig. 4G–J), we noted that in all of the extant species of *Carya* and in many species of *Juglans* we examined the internal nutshell morphology (locule configuration) corresponds to the external seed morphology, because the seed is tightly included in the nutshell (Deng & al., 2024: fig. 5). However, in extant *Juglans regia* L. (Fig. 4J) there is a gap between the inner surface of the nutshell and the ripe seed (Deng & al., 2024: fig. 7) and the internal morphology of the nutshell, without any inner symmetrical longitudinal ribs, strongly differs from that of *C. nux-taurinensis*. The extant seeds of *C. laciniosa* (F.Michx.) Loudon, *C. ovata* (Mill.) K.Koch (Fig. 4I), *C. pallida* (Ashe) Engelm. & Graebn. (Fig. 4G) and *C. tomentosa* (Lam.) Nutt. (Fig. 4H) reproduce an internal morphology of the nutshell that best matches the locule casts of *C. nux-taurinensis*, even if the external morphology is consistently smoother. Nuts of the extant *Carya aquatica* (F.Michx.) Elliott are much more similar to the fossils in external nutshell morphology, but they differ in their lateral compression and asymmetry
(Manchester, 1987: fig. 41D,E). We verified that the overall morphology of *C. nux-taurinensis* does not correspond completely to any nutshells of extant species, thus confirming its status as a distinct fossil-species.

In addition to the six above-mentioned specimens, we assign to *Carya nux-taurinensis* the following specimens from La Morra: A locule cast of a halved nut (with traces of petrified nutshell) at MNHN (MNHN.F.3010.2, "Sismonda collection"); three locule casts of complete nuts at MCCB; two almost complete (still empty) external molds of tightly closed nutshells (Fig. 3A–C; MRSN-P/345-CCN8670, 8671a, 8671b) and two locule casts of halved nuts (Fig. 3D,E; MRSN-P/345-CCN8672, 8673) sampled in 2024 (CCN). These fossils can be assigned to *C. nux-taurinensis* because they correspond in morphology to the lectotype (in particular, symmetrical longitudinal ribs of the nutshell protruding into the locule) or to the syntype. Locule casts of complete nutshells are relevant contribution of museum collections, because such a mode of preservation was not reported by Brongniart (1822) nor found during recent field observations. However, it makes sense that it was produced in the same sediment and diagenetic conditions of the other types of fossils (molds and locule casts of halved nutshells), by precipitation of minerals (likely calcite) into the fruit's seed locule after the decay of the seed and before the decay of the nutshell. One of these complete locule casts (Fig. 2B) was designated in 2023 by the Italian Palaeontological Society as the representative fossil for the Piemonte region (MCCB-04208). Another specimen assigned by us to *C. nux-taurinensis* is an external mold of the basal part of a nutshell with striking groves and three ridges deriving from post-burial mineral infilling (Fig. 3I), collected at La Morra in the eighteenth century and stored in the Allioni collection at the MGSG (Allioni, 1757; Martinetto & al., 2023). Likewise, we assign to *C. nux-taurinensis* the external mold of a probably complete nutshell (Fig. 3K) from Case Missione. There are most probably further specimens from La Morra, referable to *C. nux-taurinensis*, scattered in eighteenth and nineteenth century palaeontological collections, as we happened to detect at the MB (Fig. 3J).

Additional records of fossil fruits assigned to *Juglans nux-taurinensis*, not revised by us, concern another single site of central Italy (Gaudin & Strozzi, 1858). The assignment to *J. nux-taurinensis* of another fossil, a natural seed replica (and not nutshell!) from the surroundings of Alba (Piemonte region, NW Italy) figured in Cavallo & al. (1986: 60, fig. 76), must be rejected, because it corresponds instead to *J. regia* and probably originates from a Quaternary deposit excavated for the construction of a water well (O. Cavallo, pers. comm.). But, was the distribution of *Carya nux-taurinensis* really so limited in Europe?

Our recent discovery of new materials from the type locality La Morra clearly showed that the external and internal morphologies, together, coincide with the condition observed in organically preserved nuts previously assigned to *Carya globosa* (holotype figure reproduced by Manchester, 1987: fig. 42U,V, from the Pliocene of Dorheim, Germany, MB). This name is the result of the more recent combination of *Juglans*

*globosa* R.Ludw. (Ludwig, 1857: 103, pl. 21, figs. 12, 12a, 12b) (see Fig. 1F–H) to the genus *Carya*, validly proposed by Mädler (1939: 66, pl. 6, figs. 6–8) and accepted by, e.g., Mai (1981), Manchester (1987) and Mai & Walther (1988). These authors report *C. globosa* from the European Middle to Late Miocene and Pliocene, where it is said to be frequent (Mai & Walther, 1988). We analysed an organically preserved, almost undeformed fossil nutshell from the Pliocene of Canton Iuli (Piemonte region, NW Italy: Ucchesu & al., 2024), agreeing in the strikingly grooved external morphology (Fig. 4B,C) with *C. globosa*, and we obtained a virtual locule cast through CT-scanning (Fig. 4A,D,E). This agreed in all relevant characters to a locule cast of the complete nutshell, assigned by us to *C. nux-taurinensis*, from the Late Miocene of La Morra (Fig. 2B). We note that the merging point of the two basal lobes separated by the secondary septum is located about one-third the nut length from the base. The lectotype of *C. nux-taurinensis* shows the same merging point at half the nut length from the base, a fact which we attribute to a kind of intraspecific variability as observed in the extant fruits of *Carya*. We conclude that cast and mold fossils of La Morra (Figs. 1A–E, 2A,B, 3A–J) and the organically preserved fossils from Dorheim (Fig. 1F–H) and Canton Iuli (Fig. 4A–F) represent the same nutshell morphology.

We also assign to *Carya nux-taurinensis* the following organically preserved fossils from Italy: An unpublished, three-dimensional nutshell from the Messinian site Banengo (Martinetto & al., 2023); six nutshells, in part three-dimensional and in part compressed, from three Pliocene sites (four from Castelletto Cervo I, one from Pian Torinetto-CH10, one from Santa Barbara Gregor's outcrop: Martinetto, 2015). We do not consider here as indeed assignable to *C. nux-taurinensis* other specimens (although assigned to *C. globosa* by Martinetto, 2015: 77), because they do not show clearly such striking grooves and furrows as the nutshells from La Morra.

Although the *ICN* permits the use of separate names for fossils when the state of preservation is different, in this case such an approach appears superfluous, because the presence or absence of the organic parts does not change consistently the morphological information provided by the fossil fruits. Therefore, we consider *Carya globosa* a younger heterotypic synonym of *C. nux-taurinensis*. In this way the "Turin nuts" (Brongniart, 1822) assume increasing relevance as representatives of a rather widespread fossil-species of the genus *Carya*, extirpated from Europe in the late Cenozoic. The reliably identified fossils are sufficient to prove that the types of *C. nux-taurinensis* (Brongniart, 1822) represent the fossil record of a plant that was rather broadly distributed, not only in central Europe (Mädler, 1939; Czeczott & Skirgiełło, 1961; Mai, 1981; Manchester, 1987; Mai & Walther, 1988), but also in northern-central Italy during late Neogene (7–3 Ma).

Fossil-species:

*Carya nux-taurinensis* (Brongn.) Martinetto, De Franceschi, C.Del Rio & Manchester, **comb. nov.** [Plant Fossil
Names Registry Number: PFN003531] ≡ *Juglans nux-taurinensis* Brongn. in Mém. Mus. Hist. Nat. 8: 323, pl. VI, figs. 6a, 6b. 1822 ≡ *Juglandites nux-taurinensis* (Brongn.) Sternb., Vers. Fl. Vorwelt 1(4, Tent.): XL. 1825 – *Lectotype (designated here):* ITALY, Piemonte region, La Morra, Late Miocene, locule cast of nut-shell MNHN.F.3010.1, housed at Muséum national d'Histoire naturelle de Paris [PFN003530].

= *Carya globosa* (R.Ludw.) Mädler in Abh. Senck. Naturf. Ges. 446: 66, pl. 6, figs. 6–8. 1939 ≡ *Juglans globosa* R.Ludw. in Palaeontogr. 5: 103, pl. 21, figs. 12, 12a, 12b. 1857.

Remaining syntype of *Carex nux-taurinensis*: ITALY, Piemonte region, La Morra, Late Miocene, external mold of nutshell MNHN.F.3010.3.

## ■ AUTHOR CONTRIBUTIONS

EM studied the fossil material preserved in Italy, searched the original fossil site of *Carya nux-taurinensis* and wrote the first draft. DDF and CDR searched and studied the material at MNHN and verified and improved the draft. SRM studied the extant material, compared the morphology of extant and fossil nutshells, checked the taxonomy and finalised the manuscript together with EM. All the authors read and approved the final manuscript.

## ■ ACKNOWLEDGEMENTS

We thank Gianni Repetto for helping us to study the Italian collections and for collaboration in the field; Fabio Franceschi, Ennio Lavè, Stefano Nadile for the collaboration in the field at La Morra; Oreste Cavallo for information on the occurrence of fossil nuts in the Piemonte region; Rino Brancato and Vittorio Pane for assistance at MCCB and MGSG; Min Deng for the preparation of the virtual sections of extant nuts; Jiří Kvaček and Ludwig Luthart for the assistance in getting information from relevant old literature. Marta Moriondo largely helped us in gathering and organising data on the occasion of her bachelor thesis, defended in 2019. Open access publishing facilitated by Universita degli Studi di Torino, as part of the Wiley - CRUI-CARE agreement.

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