<?xml version="1.0"?>
<ownershipDocument>

    <schemaVersion>X0306</schemaVersion>

    <documentType>4</documentType>

    <periodOfReport>2021-08-31</periodOfReport>

    <issuer>
        <issuerCik>0001743745</issuerCik>
        <issuerName>Greenlane Holdings, Inc.</issuerName>
        <issuerTradingSymbol>GNLN</issuerTradingSymbol>
    </issuer>

    <reportingOwner>
        <reportingOwnerId>
            <rptOwnerCik>0001636985</rptOwnerCik>
            <rptOwnerName>Imbimbo Dallas</rptOwnerName>
        </reportingOwnerId>
        <reportingOwnerAddress>
            <rptOwnerStreet1>C/O GREENLANE HOLDINGS, INC.</rptOwnerStreet1>
            <rptOwnerStreet2>1095 BROKEN SOUND PARKWAY, SUITE 300</rptOwnerStreet2>
            <rptOwnerCity>BOCA RATON</rptOwnerCity>
            <rptOwnerState>FL</rptOwnerState>
            <rptOwnerZipCode>33487</rptOwnerZipCode>
            <rptOwnerStateDescription></rptOwnerStateDescription>
        </reportingOwnerAddress>
        <reportingOwnerRelationship>
            <isDirector>1</isDirector>
            <isOfficer>0</isOfficer>
            <isTenPercentOwner>0</isTenPercentOwner>
            <isOther>0</isOther>
        </reportingOwnerRelationship>
    </reportingOwner>

    <nonDerivativeTable>
        <nonDerivativeTransaction>
            <securityTitle>
                <value>Class A Common Stock</value>
            </securityTitle>
            <transactionDate>
                <value>2021-08-31</value>
            </transactionDate>
            <transactionCoding>
                <transactionFormType>4</transactionFormType>
                <transactionCode>A</transactionCode>
                <equitySwapInvolved>0</equitySwapInvolved>
            </transactionCoding>
            <transactionAmounts>
                <transactionShares>
                    <value>3021362</value>
                    <footnoteId id="F1"/>
                </transactionShares>
                <transactionPricePerShare>
                    <footnoteId id="F1"/>
                    <footnoteId id="F2"/>
                </transactionPricePerShare>
                <transactionAcquiredDisposedCode>
                    <value>A</value>
                </transactionAcquiredDisposedCode>
            </transactionAmounts>
            <postTransactionAmounts>
                <sharesOwnedFollowingTransaction>
                    <value>3021362</value>
                </sharesOwnedFollowingTransaction>
            </postTransactionAmounts>
            <ownershipNature>
                <directOrIndirectOwnership>
                    <value>D</value>
                </directOrIndirectOwnership>
            </ownershipNature>
        </nonDerivativeTransaction>
    </nonDerivativeTable>

    <derivativeTable>
        <derivativeTransaction>
            <securityTitle>
                <value>Option to Purchase</value>
            </securityTitle>
            <conversionOrExercisePrice>
                <value>4.85</value>
            </conversionOrExercisePrice>
            <transactionDate>
                <value>2021-08-31</value>
            </transactionDate>
            <transactionCoding>
                <transactionFormType>4</transactionFormType>
                <transactionCode>A</transactionCode>
                <equitySwapInvolved>0</equitySwapInvolved>
            </transactionCoding>
            <transactionAmounts>
                <transactionShares>
                    <value>28500</value>
                    <footnoteId id="F3"/>
                </transactionShares>
                <transactionPricePerShare>
                    <value>0</value>
                </transactionPricePerShare>
                <transactionAcquiredDisposedCode>
                    <value>A</value>
                </transactionAcquiredDisposedCode>
            </transactionAmounts>
            <exerciseDate>
                <footnoteId id="F3"/>
            </exerciseDate>
            <expirationDate>
                <value>2031-02-27</value>
            </expirationDate>
            <underlyingSecurity>
                <underlyingSecurityTitle>
                    <value>Class A Common Stock</value>
                </underlyingSecurityTitle>
                <underlyingSecurityShares>
                    <value>28500</value>
                </underlyingSecurityShares>
            </underlyingSecurity>
            <postTransactionAmounts>
                <sharesOwnedFollowingTransaction>
                    <value>28500</value>
                </sharesOwnedFollowingTransaction>
            </postTransactionAmounts>
            <ownershipNature>
                <directOrIndirectOwnership>
                    <value>D</value>
                </directOrIndirectOwnership>
            </ownershipNature>
        </derivativeTransaction>
        <derivativeTransaction>
            <securityTitle>
                <value>Option to Purchase</value>
            </securityTitle>
            <conversionOrExercisePrice>
                <value>2.79</value>
            </conversionOrExercisePrice>
            <transactionDate>
                <value>2021-08-31</value>
            </transactionDate>
            <transactionCoding>
                <transactionFormType>4</transactionFormType>
                <transactionCode>A</transactionCode>
                <equitySwapInvolved>0</equitySwapInvolved>
            </transactionCoding>
            <transactionAmounts>
                <transactionShares>
                    <value>28500</value>
                    <footnoteId id="F4"/>
                </transactionShares>
                <transactionPricePerShare>
                    <value>0</value>
                </transactionPricePerShare>
                <transactionAcquiredDisposedCode>
                    <value>A</value>
                </transactionAcquiredDisposedCode>
            </transactionAmounts>
            <exerciseDate>
                <footnoteId id="F4"/>
            </exerciseDate>
            <expirationDate>
                <value>2030-11-04</value>
            </expirationDate>
            <underlyingSecurity>
                <underlyingSecurityTitle>
                    <value>Class A Common Stock</value>
                </underlyingSecurityTitle>
                <underlyingSecurityShares>
                    <value>28500</value>
                </underlyingSecurityShares>
            </underlyingSecurity>
            <postTransactionAmounts>
                <sharesOwnedFollowingTransaction>
                    <value>28500</value>
                </sharesOwnedFollowingTransaction>
            </postTransactionAmounts>
            <ownershipNature>
                <directOrIndirectOwnership>
                    <value>D</value>
                </directOrIndirectOwnership>
            </ownershipNature>
        </derivativeTransaction>
        <derivativeTransaction>
            <securityTitle>
                <value>Option to Purchase</value>
            </securityTitle>
            <conversionOrExercisePrice>
                <value>2.26</value>
            </conversionOrExercisePrice>
            <transactionDate>
                <value>2021-08-31</value>
            </transactionDate>
            <transactionCoding>
                <transactionFormType>4</transactionFormType>
                <transactionCode>A</transactionCode>
                <equitySwapInvolved>0</equitySwapInvolved>
            </transactionCoding>
            <transactionAmounts>
                <transactionShares>
                    <value>3770</value>
                    <footnoteId id="F5"/>
                </transactionShares>
                <transactionPricePerShare>
                    <value>0</value>
                </transactionPricePerShare>
                <transactionAcquiredDisposedCode>
                    <value>A</value>
                </transactionAcquiredDisposedCode>
            </transactionAmounts>
            <exerciseDate>
                <footnoteId id="F5"/>
            </exerciseDate>
            <expirationDate>
                <value>2030-05-02</value>
            </expirationDate>
            <underlyingSecurity>
                <underlyingSecurityTitle>
                    <value>Class A Common Stock</value>
                </underlyingSecurityTitle>
                <underlyingSecurityShares>
                    <value>3770</value>
                </underlyingSecurityShares>
            </underlyingSecurity>
            <postTransactionAmounts>
                <sharesOwnedFollowingTransaction>
                    <value>3770</value>
                </sharesOwnedFollowingTransaction>
            </postTransactionAmounts>
            <ownershipNature>
                <directOrIndirectOwnership>
                    <value>D</value>
                </directOrIndirectOwnership>
            </ownershipNature>
        </derivativeTransaction>
        <derivativeTransaction>
            <securityTitle>
                <value>Option to Purchase</value>
            </securityTitle>
            <conversionOrExercisePrice>
                <value>2.26</value>
            </conversionOrExercisePrice>
            <transactionDate>
                <value>2021-08-31</value>
            </transactionDate>
            <transactionCoding>
                <transactionFormType>4</transactionFormType>
                <transactionCode>A</transactionCode>
                <equitySwapInvolved>0</equitySwapInvolved>
            </transactionCoding>
            <transactionAmounts>
                <transactionShares>
                    <value>3770</value>
                    <footnoteId id="F6"/>
                </transactionShares>
                <transactionPricePerShare>
                    <value>0</value>
                </transactionPricePerShare>
                <transactionAcquiredDisposedCode>
                    <value>A</value>
                </transactionAcquiredDisposedCode>
            </transactionAmounts>
            <exerciseDate>
                <footnoteId id="F6"/>
            </exerciseDate>
            <expirationDate>
                <value>2030-05-02</value>
            </expirationDate>
            <underlyingSecurity>
                <underlyingSecurityTitle>
                    <value>Class A Common Stock</value>
                </underlyingSecurityTitle>
                <underlyingSecurityShares>
                    <value>3770</value>
                </underlyingSecurityShares>
            </underlyingSecurity>
            <postTransactionAmounts>
                <sharesOwnedFollowingTransaction>
                    <value>3770</value>
                </sharesOwnedFollowingTransaction>
            </postTransactionAmounts>
            <ownershipNature>
                <directOrIndirectOwnership>
                    <value>D</value>
                </directOrIndirectOwnership>
            </ownershipNature>
        </derivativeTransaction>
        <derivativeTransaction>
            <securityTitle>
                <value>Option to Purchase</value>
            </securityTitle>
            <conversionOrExercisePrice>
                <value>1.93</value>
            </conversionOrExercisePrice>
            <transactionDate>
                <value>2021-08-31</value>
            </transactionDate>
            <transactionCoding>
                <transactionFormType>4</transactionFormType>
                <transactionCode>A</transactionCode>
                <equitySwapInvolved>0</equitySwapInvolved>
            </transactionCoding>
            <transactionAmounts>
                <transactionShares>
                    <value>24989</value>
                    <footnoteId id="F7"/>
                </transactionShares>
                <transactionPricePerShare>
                    <value>0</value>
                </transactionPricePerShare>
                <transactionAcquiredDisposedCode>
                    <value>A</value>
                </transactionAcquiredDisposedCode>
            </transactionAmounts>
            <exerciseDate>
                <footnoteId id="F7"/>
            </exerciseDate>
            <expirationDate>
                <value>2028-09-01</value>
            </expirationDate>
            <underlyingSecurity>
                <underlyingSecurityTitle>
                    <value>Class A Common Stock</value>
                </underlyingSecurityTitle>
                <underlyingSecurityShares>
                    <value>24989</value>
                </underlyingSecurityShares>
            </underlyingSecurity>
            <postTransactionAmounts>
                <sharesOwnedFollowingTransaction>
                    <value>24989</value>
                </sharesOwnedFollowingTransaction>
            </postTransactionAmounts>
            <ownershipNature>
                <directOrIndirectOwnership>
                    <value>D</value>
                </directOrIndirectOwnership>
            </ownershipNature>
        </derivativeTransaction>
        <derivativeTransaction>
            <securityTitle>
                <value>Option to Purchase</value>
            </securityTitle>
            <conversionOrExercisePrice>
                <value>1.93</value>
            </conversionOrExercisePrice>
            <transactionDate>
                <value>2021-08-31</value>
            </transactionDate>
            <transactionCoding>
                <transactionFormType>4</transactionFormType>
                <transactionCode>A</transactionCode>
                <equitySwapInvolved>0</equitySwapInvolved>
            </transactionCoding>
            <transactionAmounts>
                <transactionShares>
                    <value>24128</value>
                    <footnoteId id="F8"/>
                </transactionShares>
                <transactionPricePerShare>
                    <value>0</value>
                </transactionPricePerShare>
                <transactionAcquiredDisposedCode>
                    <value>A</value>
                </transactionAcquiredDisposedCode>
            </transactionAmounts>
            <exerciseDate>
                <footnoteId id="F8"/>
            </exerciseDate>
            <expirationDate>
                <value>2027-11-12</value>
            </expirationDate>
            <underlyingSecurity>
                <underlyingSecurityTitle>
                    <value>Class A Common Stock</value>
                </underlyingSecurityTitle>
                <underlyingSecurityShares>
                    <value>24128</value>
                </underlyingSecurityShares>
            </underlyingSecurity>
            <postTransactionAmounts>
                <sharesOwnedFollowingTransaction>
                    <value>24128</value>
                </sharesOwnedFollowingTransaction>
            </postTransactionAmounts>
            <ownershipNature>
                <directOrIndirectOwnership>
                    <value>D</value>
                </directOrIndirectOwnership>
            </ownershipNature>
        </derivativeTransaction>
    </derivativeTable>

    <footnotes>
        <footnote id="F1">Received in exchange for 10,017,779 shares of common stock of KushCo Holdings, Inc. (&quot;KushCo&quot;) pursuant to the Agreement and Plan of Merger, dated as of March 31, 2021, by and among Greenlane Holdings, Inc. (&quot;Greenlane&quot;), Merger Sub Gotham 1, LLC, Merger Sub Gotham 2, LLC and KushCo (the &quot;Merger Agreement&quot;). Upon the closing of the transactions contemplated by the Merger Agreement, each share of KushCo common stock that became fully earned and vested upon closing of the transaction contemplated by the Merger Agreement was converted into the right to receive 0.3016 shares of Greenlane Class A common stock.</footnote>
        <footnote id="F2">On the date immediately prior to the effective time of the merger, the closing price of Greenlane's Class A common stock was $2.61 and the closing price of KushCo's common stock was $0.694 per share.</footnote>
        <footnote id="F3">Received in exchange for options to purchase 94,498 shares of KushCo common stock at an exercise price of $1.46 per share pursuant to the Merger Agreement. Upon the closing of the transactions contemplated by the Merger Agreement, each option to purchase shares of KushCo common stock was converted into an option exercisable for 0.3016 shares of Greenlane Class A common stock. These options had fully vested prior to the closing of the transactions contemplated by the Merger Agreement.</footnote>
        <footnote id="F4">Received in exchange for options to purchase 94,498 shares of KushCo common stock at an exercise price of $0.84 per share pursuant to the Merger Agreement. Upon the closing of the transactions contemplated by the Merger Agreement, each option to purchase shares of KushCo common stock that became fully earned and vested upon closing of the transactions contemplated by the Merger Agreement was converted into an option exercisable for 0.3016 shares of Greenlane Class A common stock.</footnote>
        <footnote id="F5">Received in exchange for options to purchase 12,500 shares of KushCo common stock at an exercise price of $0.68 per share pursuant to the Merger Agreement. Upon the closing of the transactions contemplated by the Merger Agreement, each option to purchase shares of KushCo common stock that became fully earned and vested upon closing of the transactions contemplated by the Merger Agreement was converted into an option exercisable for 0.3016 shares of Greenlane Class A common stock.</footnote>
        <footnote id="F6">Received in exchange for options to purchase 12,500 shares of KushCo common stock at an exercise price of $0.68 per share pursuant to the Merger Agreement. Upon the closing of the transactions contemplated by the Merger Agreement, each option to purchase shares of KushCo common stock that became fully earned and vested upon closing of the transactions contemplated by the Merger Agreement was converted into an option exercisable for 0.3016 shares of Greenlane Class A common stock.</footnote>
        <footnote id="F7">Received in exchange for options to purchase 82,858 shares of KushCo common stock at an exercise price of $0.58 per share pursuant to the Merger Agreement. Upon the closing of the transactions contemplated by the Merger Agreement, each option to purchase shares of KushCo common stock was converted into an option exercisable for 0.3016 shares of Greenlane Class A common stock. These options had fully vested prior to the closing of the transactions contemplated by the Merger Agreement.</footnote>
        <footnote id="F8">Received in exchange for options to purchase 80,000 shares of KushCo common stock at an exercise price of $0.58 per share pursuant to the Merger Agreement. Upon the closing of the transactions contemplated by the Merger Agreement, each option to purchase shares of KushCo common stock that became fully earned and vested upon closing of the transactions contemplated by the Merger Agreement was converted into an option exercisable for 0.3016 shares of Greenlane Class A common stock.</footnote>
    </footnotes>

    <ownerSignature>
        <signatureName>/s/ Douglas Fischer, as attorney-in-fact for Dallas Imbimbo</signatureName>
        <signatureDate>2021-09-02</signatureDate>
    </ownerSignature>
</ownershipDocument>
